THE EFFECTS OF CROSS & SELF-FERTILISATION IN THE VEGETABLE KINGDOM.

By Charles Darwin

CONTENTS

[ DETAILED TABLE OF CONTENTS ]

[ THE EFFECTS OF CROSS AND SELF-FERTILISATION IN THE VEGETABLE KINGDOM. ]

[ CHAPTER I. INTRODUCTORY REMARKS. ]

[ CHAPTER II. CONVOLVULACEAE. ]

[ CHAPTER III. SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC. ]

[ CHAPTER IV. CRUCIFERAE, PAPAVERACEAE, RESEDACEAE, ETC. ]

[ CHAPTER V. GERANIACEAE, LEGUMINOSAE, ONAGRACEAE, ETC. ]

[ CHAPTER VI. SOLANACEAE, PRIMULACEAE, POLYGONEAE, ETC. ]

[ CHAPTER VII. A SUMMARY OF THE HEIGHTS AND WEIGHTS OF THE CROSSED AND SELF-FERTILISED PLANTS. ]

[ CHAPTER VIII. DIFFERENCE BETWEEN CROSSED AND SELF-FERTILISED PLANTS IN CONSTITUTIONAL VIGOUR AND IN OTHER RESPECTS. ]

[ CHAPTER IX. THE EFFECTS OF CROSS-FERTILISATION AND SELF-FERTILISATION ON THE PRODUCTION OF SEEDS. ]

[ CHAPTER X. MEANS OF FERTILISATION. ]

[ CHAPTER XI. THE HABITS OF INSECTS IN RELATION TO THE FERTILISATION OF FLOWERS. ]

[ CHAPTER XII. GENERAL RESULTS. ]

DETAILED TABLE OF CONTENTS

I.

INTRODUCTORY REMARKS.

Various means which favour or determine the cross-fertilisation of plants.—Benefits derived from cross-fertilisation.—Self-fertilisation favourable to the propagation of the species.—Brief history of the subject.—Object of the experiments, and the manner in which they were tried.—Statistical value of the measurements.—The experiments carried on during several successive generations.—Nature of the relationship of the plants in the later generations.—Uniformity of the conditions to which the plants were subjected.—Some apparent and some real causes of error.—Amount of pollen employed.—Arrangement of the work.—Importance of the conclusions.

II.

CONVOLVULACEAE.

Ipomoea purpurea, comparison of the height and fertility of the crossed and self-fertilised plants during ten successive generations.—Greater constitutional vigour of the crossed plants.—The effects on the offspring of crossing different flowers on the same plant, instead of crossing distinct individuals.—The effects of a cross with a fresh stock.—The descendants of the self-fertilised plant named Hero.—Summary on the growth, vigour, and fertility of the successive crossed and self-fertilised generations.—Small amount of pollen in the anthers of the self-fertilised plants of the later generations, and the sterility of their first-produced flowers.—Uniform colour of the flowers produced by the self-fertilised plants.—The advantage from a cross between two distinct plants depends on their differing in constitution.

III.

SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC.

Mimulus luteus; height, vigour, and fertility of the crossed and self-fertilised plants of the first four generations.—Appearance of a new, tall, and highly self-fertile variety.—Offspring from a cross between self-fertilised plants.—Effects of a cross with a fresh stock.—Effects of crossing flowers on the same plant.—Summary on Mimulus luteus.—Digitalis purpurea, superiority of the crossed plants.—Effects of crossing flowers on the same plant.—Calceolaria.—Linaria vulgaris.—Verbascum thapsus.—Vandellia nummularifolia.—Cleistogene flowers.—Gesneria pendulina.—Salvia coccinea.—Origanum vulgare, great increase of the crossed plants by stolons.—Thunbergia alata.

IV.

CRUCIFERAE, PAPAVERACEAE, RESEDACEAE, ETC.

Brassica oleracea, crossed and self-fertilised plants.—Great effect of a cross with a fresh stock on the weight of the offspring.—Iberis umbellata.—Papaver vagum.—Eschscholtzia californica, seedlings from a cross with a fresh stock not more vigorous, but more fertile than the self-fertilised seedlings.—Reseda lutea and odorata, many individuals sterile with their own pollen.—Viola tricolor, wonderful effects of a cross.—Adonis aestivalis.—Delphinium consolida.—Viscaria oculata, crossed plants hardly taller, but more fertile than the self-fertilised.—Dianthus caryophyllus, crossed and self-fertilised plants compared for four generations.—Great effects of a cross with a fresh stock.—Uniform colour of the flowers on the self-fertilised plants.—Hibiscus africanus.

V.

GERANIACEAE, LEGUMINOSAE, ONAGRACEAE, ETC.

Pelargonium zonale, a cross between plants propagated by cuttings does no good.—Tropaeolum minus.—Limnanthes douglasii.—Lupinus luteus and pilosus.—Phaseolus multiflorus and vulgaris.—Lathyrus odoratus, varieties of, never naturally intercross in England.—Pisum sativum, varieties of, rarely intercross, but a cross between them highly beneficial.—Sarothamnus scoparius, wonderful effects of a cross.—Ononis minutissima, cleistogene flowers of.—Summary on the Leguminosae.—Clarkia elegans.—Bartonia aurea.—Passiflora gracilis.—Apium petroselinum.—Scabiosa atropurpurea.—Lactuca sativa.—Specularia speculum.—Lobelia ramosa, advantages of a cross during two generations.—Lobelia fulgens.—Nemophila insignis, great advantages of a cross.—Borago officinalis.—Nolana prostrata.

VI.

SOLANACEAE, PRIMULACEAE, POLYGONEAE, ETC.

Petunia violacea, crossed and self-fertilised plants compared for four generations.—Effects of a cross with a fresh stock.—Uniform colour of the flowers on the self-fertilised plants of the fourth generation.—Nicotiana tabacum, crossed and self-fertilised plants of equal height.—Great effects of a cross with a distinct sub-variety on the height, but not on the fertility, of the offspring.—Cyclamen persicum, crossed seedlings greatly superior to the self-fertilised.—Anagallis collina.—Primula veris.—Equal-styled variety of Primula veris, fertility of, greatly increased by a cross with a fresh stock.—Fagopyrum esculentum.—Beta vulgaris.—Canna warscewiczi, crossed and self-fertilised plants of equal height.—Zea mays.—Phalaris canariensis.

VII.

A SUMMARY OF THE HEIGHTS AND WEIGHTS OF THE CROSSED AND SELF-FERTILISED PLANTS.

Number of species and plants measured.—Tables given.—Preliminary remarks on the offspring of plants crossed by a fresh stock.—Thirteen cases specially considered.—The effects of crossing a self-fertilised plant either by another self-fertilised plant or by an intercrossed plant of the old stock.—Summary of the results.—Preliminary remarks on the crossed and self-fertilised plants of the same stock.—The twenty-six exceptional cases considered, in which the crossed plants did not exceed greatly in height the self-fertilised.—Most of these cases shown not to be real exceptions to the rule that cross-fertilisation is beneficial.—Summary of results.—Relative weights of the crossed and self-fertilised plants.

VIII.

DIFFERENCE BETWEEN CROSSED AND SELF-FERTILISED PLANTS IN CONSTITUTIONAL VIGOUR AND IN OTHER RESPECTS.

Greater constitutional vigour of crossed plants.—The effects of great crowding.—Competition with other kinds of plants.—Self-fertilised plants more liable to premature death.—Crossed plants generally flower before the self-fertilised.—Negative effects of intercrossing flowers on the same plant.—Cases described.—Transmission of the good effects of a cross to later generations.—Effects of crossing plants of closely related parentage.—Uniform colour of the flowers on plants self-fertilised during several generations and cultivated under similar conditions.

IX.

THE EFFECTS OF CROSS-FERTILISATION AND SELF-FERTILISATION ON THE PRODUCTION OF SEEDS.

Fertility of plants of crossed and self-fertilised parentage, both lots being fertilised in the same manner.—Fertility of the parent-plants when first crossed and self-fertilised, and of their crossed and self-fertilised offspring when again crossed and self-fertilised.—Comparison of the fertility of flowers fertilised with their own pollen and with that from other flowers on the same plant.—Self-sterile plants.—Causes of self-sterility.—The appearance of highly self-fertile varieties.—Self-fertilisation apparently in some respects beneficial, independently of the assured production of seeds.—Relative weights and rates of germination of seeds from crossed and self-fertilised flowers.

X.

MEANS OF FERTILISATION.

Sterility and fertility of plants when insects are excluded.—The means by which flowers are cross-fertilised.—Structures favourable to self-fertilisation.—Relation between the structure and conspicuousness of flowers, the visits of insects, and the advantages of cross-fertilisation.—The means by which flowers are fertilised with pollen from a distinct plant.—Greater fertilising power of such pollen.—Anemophilous species.—Conversion of anemophilous species into entomophilous.—Origin of nectar.—Anemophilous plants generally have their sexes separated.—Conversion of diclinous into hermaphrodite flowers.—Trees often have their sexes separated.

XI.

THE HABITS OF INSECTS IN RELATION TO THE FERTILISATION OF FLOWERS.

Insects visit the flowers of the same species as long as they can.—Cause of this habit.—Means by which bees recognise the flowers of the same species.—Sudden secretion of nectar.—Nectar of certain flowers unattractive to certain insects.—Industry of bees, and the number of flowers visited within a short time.—Perforation of the corolla by bees.—Skill shown in the operation.—Hive-bees profit by the holes made by humble-bees.—Effects of habit.—The motive for perforating flowers to save time.—Flowers growing in crowded masses chiefly perforated.

XII.

GENERAL RESULTS.

Cross-fertilisation proved to be beneficial, and self-fertilisation injurious.—Allied species differ greatly in the means by which cross-fertilisation is favoured and self-fertilisation avoided.—The benefits and evils of the two processes depend on the degree of differentiation in the sexual elements.—The evil effects not due to the combination of morbid tendencies in the parents.—Nature of the conditions to which plants are subjected when growing near together in a state of nature or under culture, and the effects of such conditions.—Theoretical considerations with respect to the interaction of differentiated sexual elements.—Practical lessons.—Genesis of the two sexes.—Close correspondence between the effects of cross-fertilisation and self-fertilisation, and of the legitimate and illegitimate unions of heterostyled plants, in comparison with hybrid unions.

THE EFFECTS OF CROSS AND SELF-FERTILISATION IN THE VEGETABLE KINGDOM.

CHAPTER I. INTRODUCTORY REMARKS.

Various means which favour or determine the cross-fertilisation of plants.
Benefits derived from cross-fertilisation.
Self-fertilisation favourable to the propagation of the species.
Brief history of the subject.
Object of the experiments, and the manner in which they were tried.
Statistical value of the measurements.
The experiments carried on during several successive generations.
Nature of the relationship of the plants in the later generations.
Uniformity of the conditions to which the plants were subjected.
Some apparent and some real causes of error.
Amount of pollen employed.
Arrangement of the work.
Importance of the conclusions.

There is weighty and abundant evidence that the flowers of most kinds of plants are constructed so as to be occasionally or habitually cross-fertilised by pollen from another flower, produced either by the same plant, or generally, as we shall hereafter see reason to believe, by a distinct plant. Cross-fertilisation is sometimes ensured by the sexes being separated, and in a large number of cases by the pollen and stigma of the same flower being matured at different times. Such plants are called dichogamous, and have been divided into two sub-classes: proterandrous species, in which the pollen is mature before the stigma, and proterogynous species, in which the reverse occurs; this latter form of dichogamy not being nearly so common as the other. Cross-fertilisation is also ensured, in many cases, by mechanical contrivances of wonderful beauty, preventing the impregnation of the flowers by their own pollen. There is a small class of plants, which I have called dimorphic and trimorphic, but to which Hildebrand has given the more appropriate name of heterostyled; this class consists of plants presenting two or three distinct forms, adapted for reciprocal fertilisation, so that, like plants with separate sexes, they can hardly fail to be intercrossed in each generation. The male and female organs of some flowers are irritable, and the insects which touch them get dusted with pollen, which is thus transported to other flowers. Again, there is a class, in which the ovules absolutely refuse to be fertilised by pollen from the same plant, but can be fertilised by pollen from any other individual of the same species. There are also very many species which are partially sterile with their own pollen. Lastly, there is a large class in which the flowers present no apparent obstacle of any kind to self-fertilisation, nevertheless these plants are frequently intercrossed, owing to the prepotency of pollen from another individual or variety over the plant’s own pollen.

As plants are adapted by such diversified and effective means for cross-fertilisation, it might have been inferred from this fact alone that they derived some great advantage from the process; and it is the object of the present work to show the nature and importance of the benefits thus derived. There are, however, some exceptions to the rule of plants being constructed so as to allow of or to favour cross-fertilisation, for some few plants seem to be invariably self-fertilised; yet even these retain traces of having been formerly adapted for cross-fertilisation. These exceptions need not make us doubt the truth of the above rule, any more than the existence of some few plants which produce flowers, and yet never set seed, should make us doubt that flowers are adapted for the production of seed and the propagation of the species.

We should always keep in mind the obvious fact that the production of seed is the chief end of the act of fertilisation; and that this end can be gained by hermaphrodite plants with incomparably greater certainty by self-fertilisation, than by the union of the sexual elements belonging to two distinct flowers or plants. Yet it is as unmistakably plain that innumerable flowers are adapted for cross-fertilisation, as that the teeth and talons of a carnivorous animal are adapted for catching prey; or that the plumes, wings, and hooks of a seed are adapted for its dissemination. Flowers, therefore, are constructed so as to gain two objects which are, to a certain extent, antagonistic, and this explains many apparent anomalies in their structure. The close proximity of the anthers to the stigma in a multitude of species favours, and often leads, to self-fertilisation; but this end could have been gained far more safely if the flowers had been completely closed, for then the pollen would not have been injured by the rain or devoured by insects, as often happens. Moreover, in this case, a very small quantity of pollen would have been sufficient for fertilisation, instead of millions of grains being produced. But the openness of the flower and the production of a great and apparently wasteful amount of pollen are necessary for cross-fertilisation. These remarks are well illustrated by the plants called cleistogene, which bear on the same stock two kinds of flowers. The flowers of the one kind are minute and completely closed, so that they cannot possibly be crossed; but they are abundantly fertile, although producing an extremely small quantity of pollen. The flowers of the other kind produce much pollen and are open; and these can be, and often are, cross-fertilised. Hermann Muller has also made the remarkable discovery that there are some plants which exist under two forms; that is, produce on distinct stocks two kinds of hermaphrodite flowers. The one form bears small flowers constructed for self-fertilisation; whilst the other bears larger and much more conspicuous flowers plainly constructed for cross-fertilisation by the aid of insects; and without their aid these produce no seed.

The adaptation of flowers for cross-fertilisation is a subject which has interested me for the last thirty-seven years, and I have collected a large mass of observations, but these are now rendered superfluous by the many excellent works which have been lately published. In the year 1857 I wrote a short paper on the fertilisation of the kidney bean (1/1. ‘Gardeners’ Chronicle’ 1857 page 725 and 1858 pages 824 and 844. ‘Annals and Magazine of Natural History’ 3rd series volume 2 1858 page 462.); and in 1862 my work ‘On the Contrivances by which British and Foreign Orchids are Fertilised by Insects’ appeared. It seemed to me a better plan to work out one group of plants as carefully as I could, rather than to publish many miscellaneous and imperfect observations. My present work is the complement of that on Orchids, in which it was shown how admirably these plants are constructed so as to permit of, or to favour, or to necessitate cross-fertilisation. The adaptations for cross-fertilisation are perhaps more obvious in the Orchideae than in any other group of plants, but it is an error to speak of them, as some authors have done, as an exceptional case. The lever-like action of the stamens of Salvia (described by Hildebrand, Dr. W. Ogle, and others), by which the anthers are depressed and rubbed on the backs of bees, shows as perfect a structure as can be found in any orchid. Papilionaceous flowers, as described by various authors—for instance, by Mr. T.H. Farrer—offer innumerable curious adaptations for cross-fertilisation. The case of Posoqueria fragrans (one of the Rubiaceae), is as wonderful as that of the most wonderful orchid. The stamens, according to Fritz Muller, are irritable, so that as soon as a moth visits a flower, the anthers explode and cover the insect with pollen; one of the filaments which is broader than the others then moves and closes the flower for about twelve hours, after which time it resumes its original position. (1/2. ‘Botanische Zeitung’ 1866 page 129.) Thus the stigma cannot be fertilised by pollen from the same flower, but only by that brought by a moth from some other flower. Endless other beautiful contrivances for this same purpose could be specified.

Long before I had attended to the fertilisation of flowers, a remarkable book appeared in 1793 in Germany, ‘Das Entdeckte Geheimniss der Natur,’ by C.K. Sprengel, in which he clearly proved by innumerable observations, how essential a part insects play in the fertilisation of many plants. But he was in advance of his age, and his discoveries were for a long time neglected. Since the appearance of my book on Orchids, many excellent works on the fertilisation of flowers, such as those by Hildebrand, Delpino, Axell and Hermann Muller, and numerous shorter papers, have been published. (1/3. Sir John Lubbock has given an interesting summary of the whole subject in his ‘British Wild Flowers considered in relation to Insects’ 1875. Hermann Muller’s work ‘Die Befruchtung der Blumen durch Insekten’ 1873, contains an immense number of original observations and generalisations. It is, moreover, invaluable as a repertory with references to almost everything which has been published on the subject. His work differs from that of all others in specifying what kinds of insects, as far as known, visit the flowers of each species. He likewise enters on new ground, by showing not only that flowers are adapted for their own good to the visits of certain insects; but that the insects themselves are excellently adapted for procuring nectar or pollen from certain flowers. The value of H. Muller’s work can hardly be over-estimated, and it is much to be desired that it should be translated into English. Severin Axell’s work is written in Swedish, so that I have not been able to read it.) A list would occupy several pages, and this is not the proper place to give their titles, as we are not here concerned with the means, but with the results of cross-fertilisation. No one who feels interest in the mechanism by which nature effects her ends, can read these books and memoirs without the most lively interest.

From my own observations on plants, guided to a certain extent by the experience of the breeders of animals, I became convinced many years ago that it is a general law of nature that flowers are adapted to be crossed, at least occasionally, by pollen from a distinct plant. Sprengel at times foresaw this law, but only partially, for it does not appear that he was aware that there was any difference in power between pollen from the same plant and from a distinct plant. In the introduction to his book (page 4) he says, as the sexes are separated in so many flowers, and as so many other flowers are dichogamous, “it appears that nature has not willed that any one flower should be fertilised by its own pollen.” Nevertheless, he was far from keeping this conclusion always before his mind, or he did not see its full importance, as may be perceived by anyone who will read his observations carefully; and he consequently mistook the meaning of various structures. But his discoveries are so numerous and his work so excellent, that he can well afford to bear a small amount of blame. A most capable judge, H. Muller, likewise says: “It is remarkable in how very many cases Sprengel rightly perceived that pollen is necessarily transported to the stigmas of other flowers of the same species by the insects which visit them, and yet did not imagine that this transportation was of any service to the plants themselves.” (1/4. ‘Die Befruchtung der Blumen’ 1873 page 4. His words are: “Es ist merkwurdig, in wie zahlreichen Fallen Sprengel richtig erkannte, dass durch die Besuchenden Insekten der Bluthenstaub mit Nothwendigkeit auf die Narben anderer Bluthen derselben Art ubertragen wird, ohne auf die Vermuthung zu kommen, dass in dieser Wirkung der Nutzen des Insektenbesuches fur die Pflanzen selbst gesucht werden musse.”)

Andrew Knight saw the truth much more clearly, for he remarks, “Nature intended that a sexual intercourse should take place between neighbouring plants of the same species.” (1/5. ‘Philosophical Transactions’ 1799 page 202.) After alluding to the various means by which pollen is transported from flower to flower, as far as was then imperfectly known, he adds, “Nature has something more in view than that its own proper males would fecundate each blossom.” In 1811 Kolreuter plainly hinted at the same law, as did afterwards another famous hybridiser of plants, Herbert. (1/6. Kolreuter ‘Mem. de l’Acad. de St. Petersbourg’ tome 3 1809 published 1811 page 197. After showing how well the Malvaceae are adapted for cross-fertilisation, he asks, “An id aliquid in recessu habeat, quod hujuscemodi flores nunquam proprio suo pulvere, sed semper eo aliarum suae speciei impregnentur, merito quaeritur? Certe natura nil facit frustra.” Herbert ‘Amaryllidaceae, with a Treatise on Cross-bred Vegetables’ 1837.) But none of these distinguished observers appear to have been sufficiently impressed with the truth and generality of the law, so as to insist on it and impress their beliefs on others.

In 1862 I summed up my observations on Orchids by saying that nature “abhors perpetual self-fertilisation.” If the word perpetual had been omitted, the aphorism would have been false. As it stands, I believe that it is true, though perhaps rather too strongly expressed; and I should have added the self-evident proposition that the propagation of the species, whether by self-fertilisation or by cross-fertilisation, or asexually by buds, stolons, etc. is of paramount importance. Hermann Muller has done excellent service by insisting repeatedly on this latter point.

It often occurred to me that it would be advisable to try whether seedlings from cross-fertilised flowers were in any way superior to those from self-fertilised flowers. But as no instance was known with animals of any evil appearing in a single generation from the closest possible interbreeding, that is between brothers and sisters, I thought that the same rule would hold good with plants; and that it would be necessary at the sacrifice of too much time to self-fertilise and intercross plants during several successive generations, in order to arrive at any result. I ought to have reflected that such elaborate provisions favouring cross-fertilisation, as we see in innumerable plants, would not have been acquired for the sake of gaining a distant and slight advantage, or of avoiding a distant and slight evil. Moreover, the fertilisation of a flower by its own pollen corresponds to a closer form of interbreeding than is possible with ordinary bi-sexual animals; so that an earlier result might have been expected.

I was at last led to make the experiments recorded in the present volume from the following circumstance. For the sake of determining certain points with respect to inheritance, and without any thought of the effects of close interbreeding, I raised close together two large beds of self-fertilised and crossed seedlings from the same plant of Linaria vulgaris. To my surprise, the crossed plants when fully grown were plainly taller and more vigorous than the self-fertilised ones. Bees incessantly visit the flowers of this Linaria and carry pollen from one to the other; and if insects are excluded, the flowers produce extremely few seeds; so that the wild plants from which my seedlings were raised must have been intercrossed during all previous generations. It seemed therefore quite incredible that the difference between the two beds of seedlings could have been due to a single act of self-fertilisation; and I attributed the result to the self-fertilised seeds not having been well ripened, improbable as it was that all should have been in this state, or to some other accidental and inexplicable cause. During the next year, I raised for the same purpose as before two large beds close together of self-fertilised and crossed seedlings from the carnation, Dianthus caryophyllus. This plant, like the Linaria, is almost sterile if insects are excluded; and we may draw the same inference as before, namely, that the parent-plants must have been intercrossed during every or almost every previous generation. Nevertheless, the self-fertilised seedlings were plainly inferior in height and vigour to the crossed.

My attention was now thoroughly aroused, for I could hardly doubt that the difference between the two beds was due to the one set being the offspring of crossed, and the other of self-fertilised flowers. Accordingly I selected almost by hazard two other plants, which happened to be in flower in the greenhouse, namely, Mimulus luteus and Ipomoea purpurea, both of which, unlike the Linaria and Dianthus, are highly self-fertile if insects are excluded. Some flowers on a single plant of both species were fertilised with their own pollen, and others were crossed with pollen from a distinct individual; both plants being protected by a net from insects. The crossed and self-fertilised seeds thus produced were sown on opposite sides of the same pots, and treated in all respects alike; and the plants when fully grown were measured and compared. With both species, as in the cases of the Linaria and Dianthus, the crossed seedlings were conspicuously superior in height and in other ways to the self-fertilised. I therefore determined to begin a long series of experiments with various plants, and these were continued for the following eleven years; and we shall see that in a large majority of cases the crossed beat the self-fertilised plants. Several of the exceptional cases, moreover, in which the crossed plants were not victorious, can be explained.

It should be observed that I have spoken for the sake of brevity, and shall continue to do so, of crossed and self-fertilised seeds, seedlings, or plants; these terms implying that they are the product of crossed or self-fertilised flowers. Cross-fertilisation always means a cross between distinct plants which were raised from seeds and not from cuttings or buds. Self-fertilisation always implies that the flowers in question were impregnated with their own pollen.

My experiments were tried in the following manner. A single plant, if it produced a sufficiency of flowers, or two or three plants were placed under a net stretched on a frame, and large enough to cover the plant (together with the pot, when one was used) without touching it. This latter point is important, for if the flowers touch the net they may be cross-fertilised by bees, as I have known to happen; and when the net is wet the pollen may be injured. I used at first “white cotton net,” with very fine meshes, but afterwards a kind of net with meshes one-tenth of an inch in diameter; and this I found by experience effectually excluded all insects excepting Thrips, which no net will exclude. On the plants thus protected several flowers were marked, and were fertilised with their own pollen; and an equal number on the same plants, marked in a different manner, were at the same time crossed with pollen from a distinct plant. The crossed flowers were never castrated, in order to make the experiments as like as possible to what occurs under nature with plants fertilised by the aid of insects. Therefore, some of the flowers which were crossed may have failed to be thus fertilised, and afterwards have been self-fertilised. But this and some other sources of error will presently be discussed. In some few cases of spontaneously self-fertile species, the flowers were allowed to fertilise themselves under the net; and in still fewer cases uncovered plants were allowed to be freely crossed by the insects which incessantly visited them. There are some great advantages and some disadvantages in my having occasionally varied my method of proceeding; but when there was any difference in the treatment, it is always so stated under the head of each species.

Care was taken that the seeds were thoroughly ripened before being gathered. Afterwards the crossed and self-fertilised seeds were in most cases placed on damp sand on opposite sides of a glass tumbler covered by a glass plate, with a partition between the two lots; and the glass was placed on the chimney-piece in a warm room. I could thus observe the germination of the seeds. Sometimes a few would germinate on one side before any on the other, and these were thrown away. But as often as a pair germinated at the same time, they were planted on opposite sides of a pot, with a superficial partition between the two; and I thus proceeded until from half-a-dozen to a score or more seedlings of exactly the same age were planted on the opposite sides of several pots. If one of the young seedlings became sickly or was in any way injured, it was pulled up and thrown away, as well as its antagonist on the opposite side of the same pot.

As a large number of seeds were placed on the sand to germinate, many remained after the pairs had been selected, some of which were in a state of germination and others not so; and these were sown crowded together on the opposite sides of one or two rather larger pots, or sometimes in two long rows out of doors. In these cases there was the most severe struggle for life among the crossed seedlings on one side of the pot, and the self-fertilised seedlings on the other side, and between the two lots which grew in competition in the same pot. A vast number soon perished, and the tallest of the survivors on both sides when fully grown were measured. Plants treated in this manner, were subjected to nearly the same conditions as those growing in a state of nature, which have to struggle to maturity in the midst of a host of competitors.

On other occasions, from the want of time, the seeds, instead of being allowed to germinate on damp sand, were sown on the opposite sides of pots, and the fully grown plants measured. But this plan is less accurate, as the seeds sometimes germinated more quickly on one side than on the other. It was however necessary to act in this manner with some few species, as certain kinds of seeds would not germinate well when exposed to the light; though the glasses containing them were kept on the chimney-piece on one side of a room, and some way from the two windows which faced the north-east. (1/7. This occurred in the plainest manner with the seeds of Papaver vagum and Delphinium consolida, and less plainly with those of Adonis aestivalis and Ononis minutissima. Rarely more than one or two of the seeds of these four species germinated on the bare sand, though left there for some weeks; but when these same seeds were placed on earth in pots, and covered with a thin layer of sand, they germinated immediately in large numbers.)

The soil in the pots in which the seedlings were planted, or the seeds sown, was well mixed, so as to be uniform in composition. The plants on the two sides were always watered at the same time and as equally as possible; and even if this had not been done, the water would have spread almost equally to both sides, as the pots were not large. The crossed and self-fertilised plants were separated by a superficial partition, which was always kept directed towards the chief source of the light, so that the plants on both sides were equally illuminated. I do not believe it possible that two sets of plants could have been subjected to more closely similar conditions, than were my crossed and self-fertilised seedlings, as grown in the above described manner.

In comparing the two sets, the eye alone was never trusted. Generally the height of every plant on both sides was carefully measured, often more than once, namely, whilst young, sometimes again when older, and finally when fully or almost fully grown. But in some cases, which are always specified, owing to the want of time, only one or two of the tallest plants on each side were measured. This plan, which is not a good one, was never followed (except with the crowded plants raised from the seeds remaining after the pairs had been planted) unless the tallest plants on each side seemed fairly to represent the average difference between those on both sides. It has, however, some great advantages, as sickly or accidentally injured plants, or the offspring of ill-ripened seeds, are thus eliminated. When the tallest plants alone on each side were measured, their average height of course exceeds that of all the plants on the same side taken together. But in the case of the much crowded plants raised from the remaining seeds, the average height of the tallest plants was less than that of the plants in pairs, owing to the unfavourable conditions to which they were subjected from being greatly crowded. For our purpose, however, of the comparison of the crossed and self-fertilised plants, their absolute height signifies little.

As the plants were measured by an ordinary English standard divided into inches and eighths of an inch, I have not thought it worth while to change the fractions into decimals. The average or mean heights were calculated in the ordinary rough method by adding up the measurements of all, and dividing the product by the number of plants measured; the result being here given in inches and decimals. As the different species grow to various heights, I have always for the sake of easy comparison given in addition the average height of the crossed plants of each species taken as 100, and have calculated the average height of the self-fertilised plant in relation to this standard. With respect to the crowded plants raised from the seeds remaining after the pairs had been planted, and of which only some of the tallest on each side were measured, I have not thought it worth while to complicate the results by giving separate averages for them and for the pairs, but have added up all their heights, and thus obtained a single average.

I long doubted whether it was worth while to give the measurements of each separate plant, but have decided to do so, in order that it may be seen that the superiority of the crossed plants over the self-fertilised, does not commonly depend on the presence of two or three extra fine plants on the one side, or of a few very poor plants on the other side. Although several observers have insisted in general terms on the offspring from intercrossed varieties being superior to either parent-form, no precise measurements have been given (1/8. A summary of these statements, with references, may be found in my ‘Variation of Animals and Plants under Domestication’ chapter 17 2nd edition 1875 volume 2 page 109.); and I have met with no observations on the effects of crossing and self-fertilising the individuals of the same variety. Moreover, experiments of this kind require so much time—mine having been continued during eleven years—that they are not likely soon to be repeated.

As only a moderate number of crossed and self-fertilised plants were measured, it was of great importance to me to learn how far the averages were trustworthy. I therefore asked Mr. Galton, who has had much experience in statistical researches, to examine some of my tables of measurements, seven in number, namely, those of Ipomoea, Digitalis, Reseda lutea, Viola, Limnanthes, Petunia, and Zea. I may premise that if we took by chance a dozen or score of men belonging to two nations and measured them, it would I presume be very rash to form any judgment from such small numbers on their average heights. But the case is somewhat different with my crossed and self-fertilised plants, as they were of exactly the same age, were subjected from first to last to the same conditions, and were descended from the same parents. When only from two to six pairs of plants were measured, the results are manifestly of little or no value, except in so far as they confirm and are confirmed by experiments made on a larger scale with other species. I will now give the report on the seven tables of measurements, which Mr. Galton has had the great kindness to draw up for me.

[“I have examined the measurements of the plants with care, and by many statistical methods, to find out how far the means of the several sets represent constant realities, such as would come out the same so long as the general conditions of growth remained unaltered. The principal methods that were adopted are easily explained by selecting one of the shorter series of plants, say of Zea mays, for an example.”

TABLE 1/1. Zea mays (young plants). (Mr. Galton.)

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed, as recorded by Mr. Darwin.

Column 3: Self-fertilised, as recorded by Mr. Darwin.

Column 4: Crossed, in Separate Pots, arranged in order of magnitude.

Column 5: Self-fertilised, in Separate Pots, arranged in order of magnitude.

Column 6: Crossed, in a Single Series, arranged in order of magnitude.

Column 7: Self-fertilised, in a Single Series, arranged in order of magnitude.

Column 8: Difference, in a Single Series, arranged in order of magnitude.

Pot 1 : 23 4/8 : 17 3/8 :: 23 4/8 : 20 3/8 :: 23 4/8 : 20 3/8 : -3 1/8. Pot 1 : 12 : 20 3/8 :: 21 : 20 :: 23 2/8 : 20 : -3 2/8. Pot 1 : 21 : 20 :: 12 : 17 3/8 :: 23 : 20 : -3. Pot 1 : - : - :: - : - :: 22 1/8 : 18 5/8 : -3 4/8. Pot 1 : 22 : 20 :: 22 : 20 :: 22 1/8 : 18 5/8 : -3 4/8.

Pot 2 : 19 1/8 : 18 3/8 :: 21 4/8 : 18 5/8 :: 22 : 18 3/8 : -3 5/8. Pot 2 : 21 4/8 : 18 5/8 :: 19 1/8 : 18 3/8 :: 21 5/8 : 18 : -3 5/8. Pot 2 : - : - :: - : - :: 21 4/8 : 18 : -3 4/8. Pot 2 : 22 1/8 : 18 5/8 :: 23 2/8 : 18 5/8 :: 21 : 18 : -3. Pot 2 : 20 3/8 : 15 2/8 :: 22 1/8 : 18 :: 21 : 17 3/8 : -3 5/8.

Pot 3 : 18 2/8 : 16 4/8 :: 21 5/8 : 16 4/8 :: 20 3/8 : 16 4/8 : -3 7/8. Pot 3 : 21 5/8 : 18 :: 20 3/8 : 16 2/8 :: 19 1/8 : 16 2/8 : -2 7/8. Pot 3 : 23 2/8 : 16 2/8 :: 18 2/8 : 15 2/8 :: 18 2/8 : 15 4/8 : -2 6/8. Pot 3 : - : - :: - : - :: 12 : 15 2/8 : +3 2/8. Pot 3 : 21 : 18 :: 23 : 18 :: 12 : 12 6/8 : +0 6/8.

Pot 4 : 22 1/8 : 12 6/8 :: 22 1/8 : 18. Pot 4 : 23 : 15 4/8 :: 21 : 15 4/8. Pot 4 : 12 : 18 :: 12 : 12 6/8.

“The observations as I received them are shown in Table 1/1, Columns 2 and 3, where they certainly have no prima facie appearance of regularity. But as soon as we arrange them the in order of their magnitudes, as in columns 4 and 5, the case is materially altered. We now see, with few exceptions, that the largest plant on the crossed side in each pot exceeds the largest plant on the self-fertilised side, that the second exceeds the second, the third the third, and so on. Out of the fifteen cases in the table, there are only two exceptions to this rule. We may therefore confidently affirm that a crossed series will always be found to exceed a self-fertilised series, within the range of the conditions under which the present experiment has been made.”

TABLE 1/2.

Column 1: Number (Name) of Pot.

Column 2: Crossed.

Column 3: Self-fertilised.

Column 4: Difference.

Pot 1 : 18 7/8 : 19 2/8 : +0 3/8. Pot 2 : 20 7/8 : 19 : -1 7/8. Pot 3 : 21 1/8 : 16 7/8 : -4 2/8. Pot 4 : 19 6/8 : 16 : -3 6/8.

“Next as regards the numerical estimate of this excess. The mean values of the several groups are so discordant, as is shown in Table 1/2, that a fairly precise numerical estimate seems impossible. But the consideration arises, whether the difference between pot and pot may not be of much the same order of importance as that of the other conditions upon which the growth of the plants has been modified. If so, and only on that condition, it would follow that when all the measurements, either of the crossed or the self-fertilised plants, were combined into a single series, that series would be statistically regular. The experiment is tried in Table 1/1, columns 7 and 8, where the regularity is abundantly clear, and justifies us in considering its mean as perfectly reliable. I have protracted these measurements, and revised them in the usual way, by drawing a curve through them with a free hand, but the revision barely modifies the means derived from the original observations. In the present, and in nearly all the other cases, the difference between the original and revised means is under 2 per cent of their value. It is a very remarkable coincidence that in the seven kinds of plants, whose measurements I have examined, the ratio between the heights of the crossed and of the self-fertilised ranges in five cases within very narrow limits. In Zea mays it is as 100 to 84, and in the others it ranges between 100 to 76 and 100 to 86.”

“The determination of the variability (measured by what is technically called the ‘probable error’) is a problem of more delicacy than that of determining the means, and I doubt, after making many trials, whether it is possible to derive useful conclusions from these few observations. We ought to have measurements of at least fifty plants in each case, in order to be in a position to deduce fair results. One fact, however, bearing on variability, is very evident in most cases, though not in Zea mays, namely, that the self-fertilised plants include the larger number of exceptionally small specimens, while the crossed are more generally full grown.”

“Those groups of cases in which measurements have been made of a few of the tallest plants that grew in rows, each of which contained a multitude of plants, show very clearly that the crossed plants exceed the self-fertilised in height, but they do not tell by inference anything about their respective mean values. If it should happen that a series is known to follow the law of error or any other law, and if the number of individuals in the series is known, it would be always possible to reconstruct the whole series when a fragment of it has been given. But I find no such method to be applicable in the present case. The doubt as to the number of plants in each row is of minor importance; the real difficulty lies in our ignorance of the precise law followed by the series. The experience of the plants in pots does not help us to determine that law, because the observations of such plants are too few to enable us to lay down more than the middle terms of the series to which they belong with any sort of accuracy, whereas the cases we are now considering refer to one of its extremities. There are other special difficulties which need not be gone into, as the one already mentioned is a complete bar.”]

Mr. Galton sent me at the same time graphical representations which he had made of the measurements, and they evidently form fairly regular curves. He appends the words “very good” to those of Zea and Limnanthes. He also calculated the average height of the crossed and self-fertilised plants in the seven tables by a more correct method than that followed by me, namely, by including the heights, as estimated in accordance with statistical rules, of a few plants which died before they were measured; whereas I merely added up the heights of the survivors, and divided the sum by their number. The difference in our results is in one way highly satisfactory, for the average heights of the self-fertilised plants, as deduced by Mr. Galton, is less than mine in all the cases excepting one, in which our averages are the same; and this shows that I have by no means exaggerated the superiority of the crossed over the self-fertilised plants.

After the heights of the crossed and self-fertilised plants had been taken, they were sometimes cut down close to the ground, and an equal number of both weighed. This method of comparison gives very striking results, and I wish that it had been oftener followed. Finally a record was often kept of any marked difference in the rate of germination of the crossed and self-fertilised seeds,—of the relative periods of flowering of the plants raised from them,—and of their productiveness, that is, of the number of seed-capsules which they produced and of the average number of seeds which each capsule contained.

When I began my experiments I did not intend to raise crossed and self-fertilised plants for more than a single generation; but as soon as the plants of the first generation were in flower I thought that I would raise one more generation, and acted in the following manner. Several flowers on one or more of the self-fertilised plants were again self-fertilised; and several flowers on one or more of the crossed plants were fertilised with pollen from another crossed plant of the same lot. Having thus once begun, the same method was followed for as many as ten successive generations with some of the species. The seeds and seedlings were always treated in exactly the same manner as already described. The self-fertilised plants, whether originally descended from one or two mother-plants, were thus in each generation as closely interbred as was possible; and I could not have improved on my plan. But instead of crossing one of the crossed plants with another crossed plant, I ought to have crossed the self-fertilised plants of each generation with pollen taken from a non-related plant—that is, one belonging to a distinct family or stock of the same species and variety. This was done in several cases as an additional experiment, and gave very striking results. But the plan usually followed was to put into competition and compare intercrossed plants, which were almost always the offspring of more or less closely related plants, with the self-fertilised plants of each succeeding generation;—all having been grown under closely similar conditions. I have, however, learnt more by this method of proceeding, which was begun by an oversight and then necessarily followed, than if I had always crossed the self-fertilised plants of each succeeding generation with pollen from a fresh stock.

I have said that the crossed plants of the successive generations were almost always inter-related. When the flowers on an hermaphrodite plant are crossed with pollen taken from a distinct plant, the seedlings thus raised may be considered as hermaphrodite brothers or sisters; those raised from the same capsule being as close as twins or animals of the same litter. But in one sense the flowers on the same plant are distinct individuals, and as several flowers on the mother-plant were crossed by pollen taken from several flowers on the father-plant, such seedlings would be in one sense half-brothers or sisters, but more closely related than are the half-brothers and sisters of ordinary animals. The flowers on the mother-plant were, however, commonly crossed by pollen taken from two or more distinct plants; and in these cases the seedlings might be called with more truth half-brothers or sisters. When two or three mother-plants were crossed, as often happened, by pollen taken from two or three father-plants (the seeds being all intermingled), some of the seedlings of the first generation would be in no way related, whilst many others would be whole or half-brothers and sisters. In the second generation a large number of the seedlings would be what may be called whole or half first-cousins, mingled with whole and half-brothers and sisters, and with some plants not at all related. So it would be in the succeeding generations, but there would also be many cousins of the second and more remote degrees. The relationship will thus have become more and more inextricably complex in the later generations; with most of the plants in some degree and many of them closely related.

I have only one other point to notice, but this is one of the highest importance; namely, that the crossed and self-fertilised plants were subjected in the same generation to as nearly similar and uniform conditions as was possible. In the successive generations they were exposed to slightly different conditions as the seasons varied, and they were raised at different periods. But in other respects all were treated alike, being grown in pots in the same artificially prepared soil, being watered at the same time, and kept close together in the same greenhouse or hothouse. They were therefore not exposed during successive years to such great vicissitudes of climate as are plants growing out of doors.

ON SOME APPARENT AND REAL CAUSES OF ERROR IN MY EXPERIMENTS.

It has been objected to such experiments as mine, that covering plants with a net, although only for a short time whilst in flower, may affect their health and fertility. I have seen no such effect except in one instance with a Myosotis, and the covering may not then have been the real cause of injury. But even if the net were slightly injurious, and certainly it was not so in any high degree, as I could judge by the appearance of the plants and by comparing their fertility with that of neighbouring uncovered plants, it would not have vitiated my experiments; for in all the more important cases the flowers were crossed as well as self-fertilised under a net, so that they were treated in this respect exactly alike.

As it is impossible to exclude such minute pollen-carrying insects as Thrips, flowers which it was intended to fertilise with their own pollen may sometimes have been afterwards crossed with pollen brought by these insects from another flower on the same plant; but as we shall hereafter see, a cross of this kind does not produce any effect, or at most only a slight one. When two or more plants were placed near one another under the same net, as was often done, there is some real though not great danger of the flowers which were believed to be self-fertilised being afterwards crossed with pollen brought by Thrips from a distinct plant. I have said that the danger is not great because I have often found that plants which are self-sterile, unless aided by insects, remained sterile when several plants of the same species were placed under the same net. If, however, the flowers which had been presumably self-fertilised by me were in any case afterwards crossed by Thrips with pollen brought from a distinct plant, crossed seedlings would have been included amongst the self-fertilised; but it should be especially observed that this occurrence would tend to diminish and not to increase any superiority in average height, fertility, etc., of the crossed over the self-fertilised plants.

As the flowers which were crossed were never castrated, it is probable or even almost certain that I sometimes failed to cross-fertilise them effectually, and that they were afterwards spontaneously self-fertilised. This would have been most likely to occur with dichogamous species, for without much care it is not easy to perceive whether their stigmas are ready to be fertilised when the anthers open. But in all cases, as the flowers were protected from wind, rain, and the access of insects, any pollen placed by me on the stigmatic surface whilst it was immature, would generally have remained there until the stigma was mature; and the flowers would then have been crossed as was intended. Nevertheless, it is highly probable that self-fertilised seedlings have sometimes by this means got included amongst the crossed seedlings. The effect would be, as in the former case, not to exaggerate but to diminish any average superiority of the crossed over the self-fertilised plants.

Errors arising from the two causes just named, and from others,—such as some of the seeds not having been thoroughly ripened, though care was taken to avoid this error—the sickness or unperceived injury of any of the plants,—will have been to a large extent eliminated, in those cases in which many crossed and self-fertilised plants were measured and an average struck. Some of these causes of error will also have been eliminated by the seeds having been allowed to germinate on bare damp sand, and being planted in pairs; for it is not likely that ill-matured and well-matured, or diseased and healthy seeds, would germinate at exactly the same time. The same result will have been gained in the several cases in which only a few of the tallest, finest, and healthiest plants on each side of the pots were measured.

Kolreuter and Gartner have proved that with some plants several, even as many as from fifty to sixty, pollen-grains are necessary for the fertilisation of all the ovules in the ovarium. (1/9. ‘Kentniss der Befruchtung’ 1844 page 345. Naudin ‘Nouvelles Archives du Museum’ tome 1 page 27.) Naudin also found in the case of Mirabilis that if only one or two of its very large pollen-grains were placed on the stigma, the plants raised from such seeds were dwarfed. I was therefore careful to give an amply sufficient supply of pollen, and generally covered the stigma with it; but I did not take any special pains to place exactly the same amount on the stigmas of the self-fertilised and crossed flowers. After having acted in this manner during two seasons, I remembered that Gartner thought, though without any direct evidence, that an excess of pollen was perhaps injurious; and it has been proved by Spallanzani, Quatrefages, and Newport, that with various animals an excess of the seminal fluid entirely prevents fertilisation. (1/10. ‘Transactions of the Philosophical Society’ 1853 pages 253-258.) It was therefore necessary to ascertain whether the fertility of the flowers was affected by applying a rather small and an extremely large quantity of pollen to the stigma. Accordingly a very small mass of pollen-grains was placed on one side of the large stigma in sixty-four flowers of Ipomoea purpurea, and a great mass of pollen over the whole surface of the stigma in sixty-four other flowers. In order to vary the experiment, half the flowers of both lots were on plants produced from self-fertilised seeds, and the other half on plants from crossed seeds. The sixty-four flowers with an excess of pollen yielded sixty-one capsules; and excluding four capsules, each of which contained only a single poor seed, the remainder contained on an average 5.07 seeds per capsule. The sixty-four flowers with only a little pollen placed on one side of the stigma yielded sixty-three capsules, and excluding one from the same cause as before, the remainder contained on an average 5.129 seeds. So that the flowers fertilised with little pollen yielded rather more capsules and seeds than did those fertilised with an excess; but the difference is too slight to be of any significance. On the other hand, the seeds produced by the flowers with an excess of pollen were a little heavier of the two; for 170 of them weighed 79.67 grains, whilst 170 seeds from the flowers with very little pollen weighed 79.20 grains. Both lots of seeds having been placed on damp sand presented no difference in their rate of germination. We may therefore conclude that my experiments were not affected by any slight difference in the amount of pollen used; a sufficiency having been employed in all cases.

The order in which our subject will be treated in the present volume is as follows. A long series of experiments will first be given in Chapters 2 to 6. Tables will afterwards be appended, showing in a condensed form the relative heights, weights, and fertility of the offspring of the various crossed and self-fertilised species. Another table exhibits the striking results from fertilising plants, which during several generations had either been self-fertilised or had been crossed with plants kept all the time under closely similar conditions, with pollen taken from plants of a distinct stock and which had been exposed to different conditions. In the concluding chapters various related points and questions of general interest will be discussed.

Anyone not specially interested in the subject need not attempt to read all the details (though they possess, I think, some value, and cannot be all summarised. But I would suggest to the reader to take as an example the experiments on Ipomoea in Chapter 2; to which may be added those on Digitalis, Origanum, Viola, or the common cabbage, as in all these cases the crossed plants are superior to the self-fertilised in a marked degree, but not in quite the same manner. As instances of self-fertilised plants being equal or superior to the crossed, the experiments on Bartonia, Canna, and the common pea ought to be read; but in the last case, and probably in that of Canna, the want of any superiority in the crossed plants can be explained.

Species were selected for experiment belonging to widely distinct families, inhabiting various countries. In some few cases several genera belonging to the same family were tried, and these are grouped together; but the families themselves have been arranged not in any natural order, but in that which was the most convenient for my purpose. The experiments have been fully given, as the results appear to me of sufficient value to justify the details. Plants bearing hermaphrodite flowers can be interbred more closely than is possible with bisexual animals, and are therefore well-fitted to throw light on the nature and extent of the good effects of crossing, and on the evil effects of close interbreeding or self-fertilisation. The most important conclusion at which I have arrived is that the mere act of crossing by itself does no good. The good depends on the individuals which are crossed differing slightly in constitution, owing to their progenitors having been subjected during several generations to slightly different conditions, or to what we call in our ignorance spontaneous variation. This conclusion, as we shall hereafter see, is closely connected with various important physiological problems, such as the benefit derived from slight changes in the conditions of life, and this stands in the closest connection with life itself. It throws light on the origin of the two sexes and on their separation or union in the same individual, and lastly on the whole subject of hybridism, which is one of the greatest obstacles to the general acceptance and progress of the great principle of evolution.

In order to avoid misapprehension, I beg leave to repeat that throughout this volume a crossed plant, seedling, or seed, means one of crossed PARENTAGE, that is, one derived from a flower fertilised with pollen from a distinct plant of the same species. And that a self-fertilised plant, seedling, or seed, means one of self-fertilised PARENTAGE, that is, one derived from a flower fertilised with pollen from the same flower, or sometimes, when thus stated, from another flower on the same plant.

CHAPTER II. CONVOLVULACEAE.

Ipomoea purpurea, comparison of the height and fertility of the crossed
and self-fertilised plants during ten successive generations.
Greater constitutional vigour of the crossed plants.
The effects on the offspring of crossing different flowers on the same
plant, instead of crossing distinct individuals.
The effects of a cross with a fresh stock.
The descendants of the self-fertilised plant named Hero.
Summary on the growth, vigour, and fertility of the successive crossed
and self-fertilised generations.
Small amount of pollen in the anthers of the self-fertilised plants of
the later generations, and the sterility of their first-produced
flowers.
Uniform colour of the flowers produced by the self-fertilised plants.
The advantage from a cross between two distinct plants depends on their
differing in constitution.

A plant of Ipomoea purpurea, or as it is often called in England the convolvulus major, a native of South America, grew in my greenhouse. Ten flowers on this plant were fertilised with pollen from the same flower; and ten other flowers on the same plant were crossed with pollen from a distinct plant. The fertilisation of the flowers with their own pollen was superfluous, as this convolvulus is highly self-fertile; but I acted in this manner to make the experiments correspond in all respects. Whilst the flowers are young the stigma projects beyond the anthers; and it might have been thought that it could not be fertilised without the aid of humble-bees, which often visit the flowers; but as the flower grows older the stamens increase in length, and their anthers brush against the stigma, which thus receives some pollen. The number of seeds produced by the crossed and self-fertilised flowers differed very little.

[Crossed and self-fertilised seeds obtained in the above manner were allowed to germinate on damp sand, and as often as pairs germinated at the same time they were planted in the manner described in the Introduction (Chapter 1), on the opposite sides of two pots. Five pairs were thus planted; and all the remaining seeds, whether or not in a state of germination, were planted on the opposite sides of a third pot, so that the young plants on both sides were here greatly crowded and exposed to very severe competition. Rods of iron or wood of equal diameter were given to all the plants to twine up; and as soon as one of each pair reached the summit both were measured. A single rod was placed on each side of the crowded pot, Number 3, and only the tallest plant on each side was measured.

TABLE 2/1. Ipomoea purpurea (First Generation.).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Seedlings from Crossed Plants.

Column 3: Seedlings from Self-fertilised Plants.

Pot 1 : 87 4/8 : 69. Pot 1 : 87 4/8 : 66. Pot 1 : 89 : 73.

Pot 2 : 88 : 68 4/8. Pot 2 : 87 : 60 4/8.

Pot 3 : 77 : 57. Plants crowded; the tallest one measured on each side.

Total : 516 : 394.

The average height of the six crossed plants is here 86 inches, whilst that of the six self-fertilised plants is only 65.66 inches, so that the crossed plants are to the self-fertilised in height as 100 to 76. It should be observed that this difference is not due to a few of the crossed plants being extremely tall, or to a few of the self-fertilised being extremely short, but to all the crossed plants attaining a greater height than their antagonists. The three pairs in Pot 1 were measured at two earlier periods, and the difference was sometimes greater and sometimes less than that at the final measuring. But it is an interesting fact, of which I have seen several other instances, that one of the self-fertilised plants, when nearly a foot in height, was half an inch taller than the crossed plant; and again, when two feet high, it was 1 3/8 of an inch taller, but during the ten subsequent days the crossed plant began to gain on its antagonist, and ever afterward asserted its supremacy, until it exceeded its self-fertilised opponent by 16 inches.

The five crossed plants in Pots 1 and 2 were covered with a net, and produced 121 capsules; the five self-fertilised plants produced eighty-four capsules, so that the numbers of capsules were as 100 to 69. Of the 121 capsules on the crossed plants sixty-five were the product of flowers crossed with pollen from a distinct plant, and these contained on an average 5.23 seeds per capsule; the remaining fifty-six capsules were spontaneously self-fertilised. Of the eighty-four capsules on the self-fertilised plants, all the product of renewed self-fertilisation, fifty-five (which were alone examined) contained on an average 4.85 seeds per capsule. Therefore the cross-fertilised capsules, compared with the self-fertilised capsules, yielded seeds in the proportion of 100 to 93. The crossed seeds were relatively heavier than the self-fertilised seeds. Combining the above data (i.e., number of capsules and average number of contained seeds), the crossed plants, compared with the self-fertilised, yielded seeds in the ratio of 100 to 64.

These crossed plants produced, as already stated, fifty-six spontaneously self-fertilised capsules, and the self-fertilised plants produced twenty-nine such capsules. The former contained on an average, in comparison with the latter, seeds in the proportion of 100 to 99.

In Pot 3, on the opposite sides of which a large number of crossed and self-fertilised seeds had been sown and the seedlings allowed to struggle together, the crossed plants had at first no great advantage. At one time the tallest crossed was 25 1/8 inches high, and the tallest self-fertilised plants 21 3/8. But the difference afterwards became much greater. The plants on both sides, from being so crowded, were poor specimens. The flowers were allowed to fertilise themselves spontaneously under a net; the crossed plants produced thirty-seven capsules, the self-fertilised plants only eighteen, or as 100 to 47. The former contained on an average 3.62 seeds per capsule; and the latter 3.38 seeds, or as 100 to 93. Combining these data (i.e., number of capsules and average number of seeds), the crowded crossed plants produced seeds compared with the self-fertilised as 100 to 45. These latter seeds, however, were decidedly heavier, a hundred weighing 41.64 grains, than those from the capsules on the crossed plants, of which a hundred weighed 36.79 grains; and this probably was due to the fewer capsules borne by the self-fertilised plants having been better nourished. We thus see that the crossed plants in this the first generation, when grown under favourable conditions, and when grown under unfavourable conditions from being much crowded, greatly exceeded in height, and in the number of capsules produced, and slightly in the number of seeds per capsule, the self-fertilised plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

Flowers on the crossed plants of the last generation (Table 2/1) were crossed by pollen from distinct plants of the same generation; and flowers on the self-fertilised plants were fertilised by pollen from the same flower. The seeds thus produced were treated in every respect as before, and we have in Table 2/2 the result.

TABLE 2/2. Ipomoea purpurea (Second Generation.).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 87 : 67 4/8. Pot 1 : 83 : 68 4/8. Pot 1 : 83 : 80 4/8.

Pot 2 : 85 4/8 : 61 4/8. Pot 2 : 89 : 79. Pot 2 : 77 4/8 : 41.

Total : 505 : 398.

Here again every single crossed plant is taller than its antagonist. The self-fertilised plant in Pot 1, which ultimately reached the unusual height of 80 4/8 inches, was for a long time taller than the opposed crossed plant, though at last beaten by it. The average height of the six crossed plants is 84.16 inches, whilst that of the six self-fertilised plants is 66.33 inches, or as 100 to 79.

CROSSED AND SELF-FERTILISED PLANTS OF THE THIRD GENERATION.

Seeds from the crossed plants of the last generation (Table 2/2) again crossed, and from the self-fertilised plants again self-fertilised, were treated in all respects exactly as before, with the following result:—

TABLE 2/3. Ipomoea purpurea (Third Generation.).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 74 : 56 4/8. Pot 1 : 72 : 51 4/8. Pot 1 : 73 4/8 : 54.

Pot 2 : 82 : 59. Pot 2 : 81 : 30. Pot 2 : 82 : 66.

Total : 464.5 : 317.

Again all the crossed plants are higher than their antagonists: their average height is 77.41 inches, whereas that of the self-fertilised is 52.83 inches, or as 100 to 68.

I attended closely to the fertility of the plants of this third generation. Thirty flowers on the crossed plants were crossed with pollen from other crossed plants of the same generation, and the twenty-six capsules thus produced contained, on an average, 4.73 seeds; whilst thirty flowers on the self-fertilised plants, fertilised with the pollen from the same flower, produced twenty-three capsules, each containing 4.43 seeds. Thus the average number of seeds in the crossed capsules was to that in the self-fertilised capsules as 100 to 94. A hundred of the crossed seeds weighed 43.27 grains, whilst a hundred of the self-fertilised seeds weighed only 37.63 grains. Many of these lighter self-fertilised seeds placed on damp sand germinated before the crossed; thus thirty-six of the former germinated whilst only thirteen of the latter or crossed seeds germinated. In Pot 1 the three crossed plants produced spontaneously under the net (besides the twenty-six artificially cross-fertilised capsules) seventy-seven self-fertilised capsules containing on an average 4.41 seeds; whilst the three self-fertilised plants produced spontaneously (besides the twenty-three artificially self-fertilised capsules) only twenty-nine self-fertilised capsules, containing on an average 4.14 seeds. Therefore the average number of seeds in the two lots of spontaneously self-fertilised capsules was as 100 to 94. Taking into consideration the number of capsules together with the average number of seeds, the crossed plants (spontaneously self-fertilised) produced seeds in comparison with the self-fertilised plants (spontaneously self-fertilised) in the proportion of 100 to 35. By whatever method the fertility of these plants is compared, the crossed are more fertile than the self-fertilised plants.

I tried in several ways the comparative vigour and powers of growth of the crossed and self-fertilised plants of this third generation. Thus, four self-fertilised seeds which had just germinated were planted on one side of a pot, and after an interval of forty-eight hours, four crossed seeds in the same state of germination were planted on the opposite side; and the pot was kept in the hothouse. I thought that the advantage thus given to the self-fertilised seedlings would have been so great that they would never have been beaten by the crossed ones. They were not beaten until all had grown to a height of 18 inches; and the degree to which they were finally beaten is shown in Table 2/4. We here see that the average height of the four crossed plants is 76.62, and of the four self-fertilised plants 65.87 inches, or as 100 to 86; therefore less than when both sides started fair.

TABLE 2/4. Ipomoea purpurea (Third Generation, the self-fertilised plants having had a start of forty-eight hours).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 3 : 78 4/8 : 73 4/8. Pot 3 : 77 4/8 : 53. Pot 3 : 73 : 61 4/8. Pot 3 : 77 4/8 : 75 4/8.

Total : 306.5 : 263.5.

Crossed and self-fertilised seeds of the third generation were also sown out of doors late in the summer, and therefore under unfavourable conditions, and a single stick was given to each lot of plants to twine up. The two lots were sufficiently separate so as not to interfere with each other’s growth, and the ground was clear of weeds. As soon as they were killed by the first frost (and there was no difference in their hardiness), the two tallest crossed plants were found to be 24.5 and 22.5 inches, whilst the two tallest self-fertilised plants were only 15 and 12.5 inches in height, or as 100 to 59.

I likewise sowed at the same time two lots of the same seeds in a part of the garden which was shady and covered with weeds. The crossed seedlings from the first looked the most healthy, but they twined up a stick only to a height of 7 1/4 inches; whilst the self-fertilised were not able to twine at all; and the tallest of them was only 3 1/2 inches in height.

Lastly, two lots of the same seeds were sown in the midst of a bed of candy-tuft (Iberis) growing vigorously. The seedlings came up, but all the self-fertilised ones soon died excepting one, which never twined and grew to a height of only 4 inches. Many of the crossed seedlings, on the other hand, survived; and some twined up the stems of the Iberis to the height of 11 inches. These cases prove that the crossed seedlings have an immense advantage over the self-fertilised, both when growing isolated under very unfavourable conditions, and when put into competition with each other or with other plants, as would happen in a state of nature.

CROSSED AND SELF-FERTILISED PLANTS OF THE FOURTH GENERATION.

Seedlings raised as before from the crossed and self-fertilised plants of the third generation in Table 2/3, gave results as follows:—

TABLE 2/5. Ipomoea purpurea (Fourth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 84 : 80. Pot 1 : 47 : 44 1/2.

Pot 2 : 83 : 73 1/2. Pot 2 : 59 : 51 1/2.

Pot 3 : 82 : 56 1/2. Pot 3 : 65 1/2 : 63. Pot 3 : 68 : 52.

Total : 488.5 : 421.0.

Here the average height of the seven crossed plants is 69.78 inches, and that of the seven self-fertilised plants 60.14; or as 100 to 86. This smaller difference relatively to that in the former generations, may be attributed to the plants having been raised during the depth of winter, and consequently to their not having grown vigorously, as was shown by their general appearance and from several of them never reaching the summits of the rods. In Pot 2, one of the self-fertilised plants was for a long time taller by two inches than its opponent, but was ultimately beaten by it, so that all the crossed plants exceeded their opponents in height. Of twenty-eight capsules produced by the crossed plants fertilised by pollen from a distinct plant, each contained on an average 4.75 seeds; of twenty-seven self-fertilised capsules on the self-fertilised plants, each contained on an average 4.47 seeds; so that the proportion of seeds in the crossed and self-fertilised capsules was as 100 to 94.

Some of the same seeds, from which the plants in Table 2/5 had been raised, were planted, after they had germinated on damp sand, in a square tub, in which a large Brugmansia had long been growing. The soil was extremely poor and full of roots; six crossed seeds were planted in one corner, and six self-fertilised seeds in the opposite corner. All the seedlings from the latter soon died excepting one, and this grew to the height of only 1 1/2 inches. Of the crossed plants three survived, and they grew to the height of 2 1/2 inches, but were not able to twine round a stick; nevertheless, to my surprise, they produced some small miserable flowers. The crossed plants thus had a decided advantage over the self-fertilised plants under this extremity of bad conditions.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIFTH GENERATION.

These were raised in the same manner as before, and when measured gave the following results:—

TABLE 2/6. Ipomoea purpurea (Fifth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 96 : 73. Pot 1 : 86 : 78. Pot 1 : 69 : 29.

Pot 2 : 84 : 51. Pot 2 : 84 : 84. Pot 2 : 76 1/4 : 59.

Total : 495.25 : 374.00.

The average height of the six crossed plants is 82.54 inches, and that of the six self-fertilised plants 62.33 inches, or as 100 to 75. Every crossed plant exceeded its antagonist in height. In Pot 1 the middle plant on the crossed side was slightly injured whilst young by a blow, and was for a time beaten by its opponent, but ultimately recovered the usual superiority. The crossed plants produced spontaneously a vast number more capsules than did the self-fertilised plants; and the capsules of the former contained on an average 3.37 seeds, whilst those of the latter contained only 3.0 per capsule, or as 100 to 89. But looking only to the artificially fertilised capsules, those on the crossed plants again crossed contained on an average 4.46 seeds, whilst those on the self-fertilised plants again self-fertilised contained 4.77 seeds; so that the self-fertilised capsules were the more fertile of the two, and of this unusual fact I can offer no explanation.

CROSSED AND SELF-FERTILISED PLANTS OF THE SIXTH GENERATION.

These were raised in the usual manner, with the following result. I should state that there were originally eight plants on each side; but as two of the self-fertilised became extremely unhealthy and never grew to near their full height, these as well as their opponents have been struck out of the list. If they had been retained, they would have made the average height of the crossed plants unfairly greater than that of the self-fertilised. I have acted in the same manner in a few other instances, when one of a pair plainly became very unhealthy.

TABLE 2/7. Ipomoea purpurea (Sixth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 93 : 50 1/2. Pot 1 : 91 : 65.

Pot 2 : 79 : 50. Pot 2 : 86 1/2 : 87. Pot 2 : 88 : 62.

Pot 3 : 87 1/2 : 64 1/2.

Total : 525 : 379.

The average height of the six crossed plants is here 87.5, and of the six self-fertilised plants 63.16, or as 100 to 72. This large difference was chiefly due to most of the plants, especially the self-fertilised ones, having become unhealthy towards the close of their growth, and they were severely attacked by aphides. From this cause nothing can be inferred with respect to their relative fertility. In this generation we have the first instance of a self-fertilised plant in Pot 2 exceeding (though only by half an inch) its crossed opponent. This victory was fairly won after a long struggle. At first the self-fertilised plant was several inches taller than its opponent, but when the latter was 4 1/2 feet high it had grown equal; it then grew a little taller than the self-fertilised plant, but was ultimately beaten by it to the extent of half an inch, as shown in Table 2/7. I was so much surprised at this case that I saved the self-fertilised seeds of this plant, which I will call the “Hero,” and experimented on its descendants, as will hereafter be described.

Besides the plants included in Table 2/7, nine crossed and nine self-fertilised plants of the same lot were raised in two other pots, 4 and 5. These pots had been kept in the hothouse, but from want of room were, whilst the plants were young, suddenly moved during very cold weather into the coldest part of the greenhouse. They all suffered greatly, and never quite recovered. After a fortnight only two of the nine self-fertilised seedlings were alive, whilst seven of the crossed survived. The tallest of these latter plants when measured was 47 inches in height, whilst the tallest of the two surviving self-fertilised plants was only 32 inches. Here again we see how much more vigorous the crossed plants are than the self-fertilised.

CROSSED AND SELF-FERTILISED PLANTS OF THE SEVENTH GENERATION.

These were raised as heretofore with the following result:—

TABLE 2/8. Ipomoea purpurea (Seventh Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 84 4/8 : 74 6/8. Pot 1 : 84 6/8 : 84. Pot 1 : 76 2/8 : 55 4/8.

Pot 2 : 84 4/8 : 65. Pot 2 : 90 : 51 2/8. Pot 2 : 82 2/8 : 80 4/8.

Pot 3 : 83 : 67 6/8. Pot 3 : 86 : 60 2/8.

Pot 4 : 84 2/8 : 75 2/8.

Total : 755.50 : 614.25.

Each of these nine crossed plants is higher than its opponent, though in one case only by three-quarters of an inch. Their average height is 83.94 inches, and that of the self-fertilised plants 68.25, or as 100 to 81. These plants, after growing to their full height, became very unhealthy and infested with aphides, just when the seeds were setting, so that many of the capsules failed, and nothing can be said on their relative fertility.

CROSSED AND SELF-FERTILISED PLANTS OF THE EIGHTH GENERATION.

As just stated, the plants of the last generation, from which the present ones were raised, were very unhealthy and their seeds of unusually small size; and this probably accounts for the two lots behaving differently to what they did in any of the previous or succeeding generations. Many of the self-fertilised seeds germinated before the crossed ones, and these were of course rejected. When the crossed seedlings in Table 2/9 had grown to a height of between 1 and 2 feet, they were all, or almost all, shorter than their self-fertilised opponents, but were not then measured. When they had acquired an average height of 32.28 inches, that of the self-fertilised plants was 40.68, or as 100 to 122. Moreover, every one of the self-fertilised plants, with a single exception, exceeded its crossed opponent. When, however, the crossed plants had grown to an average height of 77.56 inches, they just exceeded (namely, by .7 of an inch) the average height of the self-fertilised plants; but two of the latter were still taller than their crossed opponents. I was so much astonished at this whole case, that I tied string to the summits of the rods; the plants being thus allowed to continue climbing upwards. When their growth was complete they were untwined, stretched straight, and measured. The crossed plants had now almost regained their accustomed superiority, as may be seen in Table 2/9.

The average height of the eight crossed plants is here 113.25 inches, and that of the self-fertilised plants 96.65, or as 100 to 85. Nevertheless two of the self-fertilised plants, as may be seen in Table 2/9, were still higher than their crossed opponents. The latter manifestly had much thicker stems and many more lateral branches, and looked altogether more vigorous than the self-fertilised plants, and generally flowered before them. The earlier flowers produced by these self-fertilised plants did not set any capsules, and their anthers contained only a small amount of pollen; but to this subject I shall return. Nevertheless capsules produced by two other self-fertilised plants of the same lot, not included in Table 2/9, which had been highly favoured by being grown in separate pots, contained the large average number of 5.1 seeds per capsule.

TABLE 2/9. Ipomoea purpurea (Eighth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 111 6/8 : 96. Pot 1 : 127 : 54. Pot 1 : 130 6/8 : 93 4/8.

Pot 2 : 97 2/8 : 94. Pot 2 : 89 4/8 : 125 6/8.

Pot 3 : 103 6/8 : 115 4/8. Pot 3 : 100 6/8 : 84 6/8. Pot 3 : 147 4/8 : 109 6/8.

Total : 908.25 : 773.25.

CROSSED AND SELF-FERTILISED PLANTS OF THE NINTH GENERATION.

The plants of this generation were raised in the same manner as before, with the result shown in Table 2/10.

The fourteen crossed plants average in height 81.39 inches and the fourteen self-fertilised plants 64.07, or as 100 to 79. One self-fertilised plant in Pot 3 exceeded, and one in Pot 4 equalled in height, its opponent. The self-fertilised plants showed no sign of inheriting the precocious growth of their parents; this having been due, as it would appear, to the abnormal state of the seeds from the unhealthiness of their parents. The fourteen self-fertilised plants yielded only forty spontaneously self-fertilised capsules, to which must be added seven, the product of ten flowers artificially self-fertilised. On the other hand, the fourteen crossed plants yielded 152 spontaneously self-fertilised capsules; but thirty-six flowers on these plants were crossed (yielding thirty-three capsules), and these flowers would probably have produced about thirty spontaneously self-fertilised capsules. Therefore an equal number of the crossed and self-fertilised plants would have produced capsules in the proportion of about 182 to 47, or as 100 to 26. Another phenomenon was well pronounced in this generation, but I believe had occurred previously to a slight extent; namely, that most of the flowers on the self-fertilised plants were somewhat monstrous. The monstrosity consisted in the corolla being irregularly split so that it did not open properly, with one or two of the stamens slightly foliaceous, coloured, and firmly coherent to the corolla. I observed this monstrosity in only one flower on the crossed plants. The self-fertilised plants, if well nourished, would almost certainly, in a few more generations, have produced double flowers, for they had already become in some degree sterile. (2/1. See on this subject ‘Variation of Animals and Plants under Domestication’ chapter 18 2nd edition volume 2 page 152.)

TABLE 2/10. Ipomoea purpurea (Ninth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 83 4/8 : 57. Pot 1 : 85 4/8 : 71. Pot 1 : 83 4/8 : 48 3/8.

Pot 2 : 83 2/8 : 45. Pot 2 : 64 2/8 : 43 6/8. Pot 2 : 64 3/8 : 38 4/8.

Pot 3 : 79 : 63. Pot 3 : 88 1/8 : 71. Pot 3 : 61 : 89 4/8.

Pot 4 : 82 4/8 : 82 4/8. Pot 4 : 90 : 76 1/8.

Pot 5 : 89 4/8 : 67. Pot 5 : 92 4/8 : 74 2/8. Pot 5 : 92 4/8 : 70. Crowded plants.

Total : 1139.5 : 897.0.

CROSSED AND SELF-FERTILISED PLANTS OF THE TENTH GENERATION.

Six plants were raised in the usual manner from the crossed plants of the last generation (Table 2/10) again intercrossed, and from the self-fertilised again self-fertilised. As one of the crossed plants in Pot 1 in Table 2/11 became much diseased, having crumpled leaves, and producing hardly any capsules, it and its opponent have been struck out of the table.

TABLE 2/11. Ipomoea purpurea (Tenth Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 92 3/8 : 47 2/8. Pot 1 : 94 4/8 : 34 6/8.

Pot 2 : 87 : 54 4/8. Pot 2 : 89 5/8 : 49 2/8. Pot 2 : 105 : 66 2/8.

Total : 468.5 : 252.0.

The five crossed plants average 93.7 inches, and the five self-fertilised only 50.4, or as 100 to 54. This difference, however, is so great that it must be looked at as in part accidental. The six crossed plants (the diseased one here included) yielded spontaneously 101 capsules, and the six self-fertilised plants 88, the latter being chiefly produced by one of the plants. But as the diseased plant, which yielded hardly any seed, is here included, the ratio of 101 to 88 does not fairly give the relative fertility of the two lots. The stems of the six crossed plants looked so much finer than those of the six self-fertilised plants, that after the capsules had been gathered and most of the leaves had fallen off, they were weighed. Those of the crossed plants weighed 2,693 grains, whilst those of the self-fertilised plants weighed only 1,173 grains, or as 100 to 44; but as the diseased and dwarfed crossed plant is here included, the superiority of the former in weight was really greater.]

THE EFFECTS ON THE OFFSPRING OF CROSSING DIFFERENT FLOWERS ON THE SAME PLANT, INSTEAD OF CROSSING DISTINCT INDIVIDUALS.

In all the foregoing experiments, seedlings from flowers crossed by pollen from a distinct plant (though in the later generations more or less closely related) were put into competition with, and almost invariably proved markedly superior in height to the offspring from self-fertilised flowers. I wished, therefore, to ascertain whether a cross between two flowers on the same plant would give to the offspring any superiority over the offspring from flowers fertilised with their own pollen. I procured some fresh seed and raised two plants, which were covered with a net; and several of their flowers were crossed with pollen from a distinct flower on the same plant. Twenty-nine capsules thus produced contained on an average 4.86 seeds per capsule; and 100 of these seeds weighed 36.77 grains. Several other flowers were fertilised with their own pollen, and twenty-six capsules thus produced contained on an average 4.42 seeds per capsule; 100 of which weighed 42.61 grains. So that a cross of this kind appears to have increased slightly the number of seeds per capsule, in the ratio of 100 to 91; but these crossed seeds were lighter than the self-fertilised in the ratio of 86 to 100. I doubt, however, from other observations, whether these results are fully trustworthy. The two lots of seeds, after germinating on sand, were planted in pairs on the opposite sides of nine pots, and were treated in every respect like the plants in the previous experiments. The remaining seeds, some in a state of germination and some not so, were sown on the opposite sides of a large pot (Number 10); and the four tallest plants on each side of this pot were measured. The result is shown in Table 2/12.

TABLE 2/12. Ipomoea purpurea.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 82 : 77 4/8. Pot 1 : 75 : 87. Pot 1 : 65 : 64. Pot 1 : 76 : 87 2/8.

Pot 2 : 78 4/8 : 84. Pot 2 : 43 : 86 4/8. Pot 2 : 65 4/8 : 90 4/8.

Pot 3 : 61 2/8 : 86. Pot 3 : 85 : 69 4/8. Pot 3 : 89 : 87 4/8.

Pot 4 : 83 : 80 4/8. Pot 4 : 73 4/8 : 88 4/8. Pot 4 : 67 : 84 4/8.

Pot 5 : 78 : 66 4/8. Pot 5 : 76 6/8 : 77 4/8. Pot 5 : 57 : 81 4/8.

Pot 6 : 70 4/8 : 80. Pot 6 : 79 : 82 4/8. Pot 6 : 79 6/8 : 55 4/8.

Pot 7 : 76 : 77. Pot 7 : 84 4/8 : 83 4/8. Pot 7 : 79 : 73 4/8.

Pot 8 : 73 : 76 4/8. Pot 8 : 67 : 82. Pot 8 : 83 : 80 4/8.

Pot 9 : 73 2/8 : 78 4/8. Pot 9 : 78 : 67 4/8.

Pot 10 : 34 : 82 4/8. Pot 10 : 82 : 36 6/8. Pot 10 : 84 6/8 : 69 4/8. Pot 10 : 71 : 75 2/8. Crowded plants.

Total : 2270.25 : 2399.75.

The average height of the thirty-one crossed plants is 73.23 inches, and that of the thirty-one self-fertilised plants 77.41 inches; or as 100 to 106. Looking to each pair, it may be seen that only thirteen of the crossed plants, whilst eighteen of the self-fertilised plants exceed their opponents. A record was kept with respect to the plant which flowered first in each pot; and only two of the crossed flowered before one of the self-fertilised in the same pot; whilst eight of the self-fertilised flowered first. It thus appears that the crossed plants are slightly inferior in height and in earliness of flowering to the self-fertilised. But the inferiority in height is so small, namely as 100 to 106, that I should have felt very doubtful on this head, had I not cut down all the plants (except those in the crowded pot Number 10) close to the ground and weighed them. The twenty-seven crossed plants weighed 16 1/2 ounces, and the twenty-seven self-fertilised plants 20 1/2 ounces; and this gives a ratio of 100 to 124.

A self-fertilised plant of the same parentage as those in Table 2/12 had been raised in a separate pot for a distinct purpose; and it proved partially sterile, the anthers containing very little pollen. Several flowers on this plant were crossed with the little pollen which could be obtained from the other flowers on the same plant; and other flowers were self-fertilised. From the seeds thus produced four crossed and four self-fertilised plants were raised, which were planted in the usual manner on the opposite sides of two pots. All these four crossed plants were inferior in height to their opponents; they averaged 78.18 inches, whilst the four self-fertilised plants averaged 84.8 inches; or as 100 to 108. (2/2. From one of these self-fertilised plants, spontaneously self-fertilised, I gathered twenty-four capsules, and they contained on an average only 3.2 seeds per capsule; so that this plant had apparently inherited some of the sterility of its parent.) This case, therefore, confirms the last. Taking all the evidence together, we must conclude that these strictly self-fertilised plants grew a little taller, were heavier, and generally flowered before those derived from a cross between two flowers on the same plant. These latter plants thus present a wonderful contrast with those derived from a cross between two distinct individuals.

THE EFFECTS ON THE OFFSPRING OF A CROSS WITH A DISTINCT OR FRESH STOCK BELONGING TO THE SAME VARIETY.

From the two foregoing series of experiments we see, firstly, the good effects during several successive generations of a cross between distinct plants, although these were in some degree inter-related and had been grown under nearly the same conditions; and, secondly, the absence of all such good effects from a cross between flowers on the same plant; the comparison in both cases being made with the offspring of flowers fertilised with their own pollen. The experiments now to be given show how powerfully and beneficially plants, which have been intercrossed during many successive generations, having been kept all the time under nearly uniform conditions, are affected by a cross with another plant belonging to the same variety, but to a distinct family or stock, which had grown under different conditions.

[Several flowers on the crossed plants of the ninth generation in Table 2/10, were crossed with pollen from another crossed plant of the same lot. The seedlings thus raised formed the tenth intercrossed generation, and I will call them the “INTERCROSSED PLANTS.” Several other flowers on the same crossed plants of the ninth generation were fertilised (not having been castrated) with pollen taken from plants of the same variety, but belonging to a distinct family, which had been grown in a distant garden at Colchester, and therefore under somewhat different conditions. The capsules produced by this cross contained, to my surprise, fewer and lighter seeds than did the capsules of the intercrossed plants; but this, I think, must have been accidental. The seedlings raised from them I will call the “COLCHESTER-CROSSED.” The two lots of seeds, after germinating on sand, were planted in the usual manner on the opposite sides of five pots, and the remaining seeds, whether or not in a state of germination, were thickly sown on the opposite sides of a very large pot, Number 6 in Table 2/13. In three of the six pots, after the young plants had twined a short way up their sticks, one of the Colchester-crossed plants was much taller than any one of the intercrossed plants on the opposite side of the same pot; and in the three other pots somewhat taller. I should state that two of the Colchester-crossed plants in Pot 4, when about two-thirds grown, became much diseased, and were, together with their intercrossed opponents, rejected. The remaining nineteen plants, when almost fully grown, were measured, with the following result:

TABLE 2/13. Ipomoea purpurea.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Colchester-Crossed Plants.

Column 3: Intercrossed Plants of the Tenth Generation.

Pot 1 : 87 : 78. Pot 1 : 87 4/8 : 68 4/8. Pot 1 : 85 1/8 : 94 4/8.

Pot 2 : 93 6/8 : 60. Pot 2 : 85 4/8 : 87 2/8. Pot 2 : 90 5/8 : 45 4/8.

Pot 3 : 84 2/8 : 70 1/8. Pot 3 : 92 4/8 : 81 6/8. Pot 3 : 85 : 86 2/8.

Pot 4 : 95 6/8 : 65 1/8.

Pot 5 : 90 4/8 : 85 6/8. Pot 5 : 86 6/8 : 63. Pot 5 : 84 : 62 6/8.

Pot 6 : 90 4/8 : 43 4/8. Pot 6 : 75 : 39 6/8. Pot 6 : 71 : 30 2/8. Pot 6 : 83 6/8 : 86. Pot 6 : 63 : 53. Pot 6 : 65 : 48 6/8. Crowded plants in a very large pot.

Total : 1596.50 : 1249.75.

In sixteen out of these nineteen pairs, the Colchester-crossed plant exceeded in height its intercrossed opponent. The average height of the Colchester-crossed is 84.03 inches, and that of the intercrossed 65.78 inches; or as 100 to 78. With respect to the fertility of the two lots, it was too troublesome to collect and count the capsules on all the plants; so I selected two of the best pots, 5 and 6, and in these the Colchester-crossed produced 269 mature and half-mature capsules, whilst an equal number of the intercrossed plants produced only 154 capsules; or as 100 to 57. By weight the capsules from the Colchester-crossed plants were to those from the intercrossed plants as 100 to 51; so that the former probably contained a somewhat larger average number of seeds.]

We learn from this important experiment that plants in some degree related, which had been intercrossed during the nine previous generations, when they were fertilised with pollen from a fresh stock, yielded seedlings as superior to the seedlings of the tenth intercrossed generation, as these latter were to the self-fertilised plants of the corresponding generation. For if we look to the plants of the ninth generation in Table 2/10 (and these offer in most respects the fairest standard of comparison) we find that the intercrossed plants were in height to the self-fertilised as 100 to 79, and in fertility as 100 to 26; whilst the Colchester-crossed plants are in height to the intercrossed as 100 to 78, and in fertility as 100 to 51.

[THE DESCENDANTS OF THE SELF-FERTILISED PLANT, NAMED HERO, WHICH APPEARED IN THE SIXTH SELF-FERTILISED GENERATION.

In the five generations before the sixth, the crossed plant of each pair was taller than its self-fertilised opponent; but in the sixth generation (Table 2/7, Pot 2) the Hero appeared, which after a long and dubious struggle conquered its crossed opponent, though by only half an inch. I was so much surprised at this fact, that I resolved to ascertain whether this plant would transmit its powers of growth to its seedlings. Several flowers on Hero were therefore fertilised with their own pollen, and the seedlings thus raised were put into competition with self-fertilised and intercrossed plants of the corresponding generation. The three lots of seedlings thus all belong to the seventh generation. Their relative heights are shown in Tables 2/14 and 2/15.

TABLE 2/14. Ipomoea purpurea.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Self-fertilised Plants of the Seventh Generation, Children of Hero.

Column 3: Self-fertilised Plants of the Seventh Generation.

Pot 1 : 74 : 89 4/8. Pot 1 : 60 : 61. Pot 1 : 55 2/8 : 49.

Pot 2 : 92 : 82. Pot 2 : 91 6/8 : 56. Pot 2 : 74 2/8 : 38.

Total : 447.25 : 375.50.

The average height of the six self-fertilised children of Hero is 74.54 inches, whilst that of the ordinary self-fertilised plants of the corresponding generation is only 62.58 inches, or as 100 to 84.

TABLE 2/15. Ipomoea purpurea.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Self-fertilised Plants of the Seventh Generation, Children of Hero.

Column 3: Intercrossed Plants of the Seventh Generation.

Pot 3 : 92 : 76 6/8.

Pot 4 : 87 : 89. Pot 4 : 87 6/8 : 86 6/8.

Total : 266.75 : 252.50.

Here the average height of the three self-fertilised children of Hero is 88.91 inches, whilst that of the intercrossed plants is 84.16; or as 100 to 95. We thus see that the self-fertilised children of Hero certainly inherit the powers of growth of their parents; for they greatly exceed in height the self-fertilised offspring of the other self-fertilised plants, and even exceed by a trifle the intercrossed plants,—all of the corresponding generation.

Several flowers on the self-fertilised children of Hero in Table 2/14 were fertilised with pollen from the same flower; and from the seeds thus produced, self-fertilised plants of the eighth generation (grandchildren of Hero) were raised. Several other flowers on the same plants were crossed with pollen from the other children of Hero. The seedlings raised from this cross may be considered as the offspring of the union of brothers and sisters. The result of the competition between these two sets of seedlings (namely self-fertilised and the offspring of brothers and sisters) is given in Table 2/16.

TABLE 2/16. Ipomoea purpurea.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Self-fertilised Grandchildren of Hero, from the Self-fertilised Children. Eighth Generation.

Column 3: Grandchildren from a cross between the self-fertilised children of Hero. Eighth Generation.

Pot 1 : 86 6/8 : 95 6/8. Pot 1 : 90 3/8 : 95 3/8.

Pot 2 : 96 : 85. Pot 2 : 77 2/8 : 93.

Pot 3 : 73 : 86 2/8. Pot 3 : 66 : 82 2/8. Pot 3 : 84 4/8 : 70 6/8.

Pot 4 : 88 1/8 : 66 3/8. Pot 4 : 84 : 15 4/8. Pot 4 : 36 2/8 : 38. Pot 4 : 74 : 78 3/8.

Pot 5 : 90 1/8 : 82 6/8. Pot 5 : 90 5/8 : 83 6/8.

Total : 1037.00 : 973.16.

The average height of the thirteen self-fertilised grandchildren of Hero is 79.76 inches, and that of the grandchildren from a cross between the self-fertilised children is 74.85; or as 100 to 94. But in Pot 4 one of the crossed plants grew only to a height of 15 1/2 inches; and if this plant and its opponent are struck out, as would be the fairest plan, the average height of the crossed plants exceeds only by a fraction of an inch that of the self-fertilised plants. It is therefore clear that a cross between the self-fertilised children of Hero did not produce any beneficial effect worth notice; and it is very doubtful whether this negative result can be attributed merely to the fact of brothers and sisters having been united, for the ordinary intercrossed plants of the several successive generations must often have been derived from the union of brothers and sisters (as shown in Chapter 1), and yet all of them were greatly superior to the self-fertilised plants. We are therefore driven to the suspicion, which we shall soon see strengthened, that Hero transmitted to its offspring a peculiar constitution adapted for self-fertilisation.

It would appear that the self-fertilised descendants of Hero have not only inherited from Hero a power of growth equal to that of the ordinary intercrossed plants, but have become more fertile when self-fertilised than is usual with the plants of the present species. The flowers on the self-fertilised grandchildren of Hero in Table 2.16 (the eighth generation of self-fertilised plants) were fertilised with their own pollen and produced plenty of capsules, ten of which (though this is too few a number for a safe average) contained 5.2 seeds per capsule,—a higher average than was observed in any other case with the self-fertilised plants. The anthers produced by these self-fertilised grandchildren were also as well developed and contained as much pollen as those on the intercrossed plants of the corresponding generation; whereas this was not the case with the ordinary self-fertilised plants of the later generations. Nevertheless some few of the flowers produced by the grandchildren of Hero were slightly monstrous, like those of the ordinary self-fertilised plants of the later generations. In order not to recur to the subject of fertility, I may add that twenty-one self-fertilised capsules, spontaneously produced by the great-grandchildren of Hero (forming the ninth generation of self-fertilised plants), contained on an average 4.47 seeds; and this is as high an average as the self-fertilised flowers of any generation usually yielded.

Several flowers on the self-fertilised grandchildren of Hero in Table 2/16 were fertilised with pollen from the same flower; and the seedlings raised from them (great-grandchildren of Hero) formed the ninth self-fertilised generation. Several other flowers were crossed with pollen from another grandchild, so that they may be considered as the offspring of brothers and sisters, and the seedlings thus raised may be called the INTERCROSSED great-grandchildren. And lastly, other flowers were fertilised with pollen from a distinct stock, and the seedlings thus raised may be called the COLCHESTER-CROSSED great-grandchildren. In my anxiety to see what the result would be, I unfortunately planted the three lots of seeds (after they had germinated on sand) in the hothouse in the middle of winter, and in consequence of this the seedlings (twenty in number of each kind) became very unhealthy, some growing only a few inches in height, and very few to their proper height. The result, therefore, cannot be fully trusted; and it would be useless to give the measurements in detail. In order to strike as fair an average as possible, I first excluded all the plants under 50 inches in height, thus rejecting all the most unhealthy plants. The six self-fertilised thus left were on an average 66.86 inches high; the eight intercrossed plants 63.2 high; and the seven Colchester-crossed 65.37 high; so that there was not much difference between the three sets, the self-fertilised plants having a slight advantage. Nor was there any great difference when only the plants under 36 inches in height were excluded. Nor again when all the plants, however much dwarfed and unhealthy, were included. In this latter case the Colchester-crossed gave the lowest average of all; and if these plants had been in any marked manner superior to the other two lots, as from my former experience I fully expected they would have been, I cannot but think that some vestige of such superiority would have been evident, notwithstanding the very unhealthy condition of most of the plants. No advantage, as far as we can judge, was derived from intercrossing two of the grandchildren of Hero, any more than when two of the children were crossed. It appears therefore that Hero and its descendants have varied from the common type, not only in acquiring great power of growth, and increased fertility when subjected to self-fertilisation, but in not profiting from a cross with a distinct stock; and this latter fact, if trustworthy, is a unique case, as far as I have observed in all my experiments.]

A SUMMARY ON THE GROWTH, VIGOUR, AND FERTILITY OF THE SUCCESSIVE GENERATIONS OF THE CROSSED AND SELF-FERTILISED PLANTS OF Ipomoea purpurea, TOGETHER WITH SOME MISCELLANEOUS OBSERVATIONS.

In Table 2/17, we see the average or mean heights of the ten successive generations of the intercrossed and self-fertilised plants, grown in competition with each other; and in the right hand column we have the ratios of the one to the other, the height of the intercrossed plants being taken at 100. In the bottom line the mean height of the seventy-three intercrossed plants is shown to be 85.84 inches, and that of the seventy-three self-fertilised plants 66.02 inches, or as 100 to 77.

TABLE 2/17. Ipomoea purpurea. Summary of measurements of the ten generations.

Heights of Plants in inches:

Column 1: Name of Generation.

Column 2: Number of Crossed Plants.

Column 3: Average Height of Crossed Plants.

Column 4: Number of Self-fertilised Plants.

Column 5: Average Height of Self-fertilised Plants.

Column 6: n in Ratio between Average Heights of Crossed and Self-fertilised Plants, expressed as 100 to n.

First generation Table 2/1 : 6 : 86.00 : 6 : 65.66 : 76.

Second generation Table 2/2 : 6 : 84.16 : 6 : 66.33 : 79.

Third generation Table 2/3 : 6 : 77.41 : 6 : 52.83 : 68.

Fourth generation Table 2/5 : 7 : 69.78 : 7 : 60.14 : 86.

Fifth generation Table 2/6 : 6 : 82.54 : 6 : 62.33 : 75.

Sixth generation Table 2/7 : 6 : 87.50 : 6 : 63.16 : 72.

Seventh generation Table 2/8 : 9 : 83.94 : 9 : 68.25 : 81.

Eighth generation Table 2/9 : 8 : 113.25 : 8 : 96.65 : 85.

Ninth generation Table 2/10 : 14 : 81.39 : 14 : 64.07 : 79.

Tenth generation Table 2/11 : 5 : 93.70 : 5 : 50.40 : 54.

All ten generations together : 73 : 85.84 : 73 : 66.02 : 77.

(DIAGRAM 2/1. Diagram showing the mean heights of the crossed and self-fertilised plants of Ipomoea purpurea in the ten generations; the mean height of the crossed plants being taken as 100. On the right hand, the mean heights of the crossed and self-fertilised plants of all the generations taken together are shown (as eleven pairs of unequal vertical lines.))

The mean height of the self-fertilised plants in each of the ten generations is also shown in the diagram 2/1, that of the intercrossed plants being taken at 100, and on the right side we see the relative heights of the seventy-three intercrossed plants, and of the seventy-three self-fertilised plants. The difference in height between the crossed and self-fertilised plants will perhaps be best appreciated by an illustration: If all the men in a country were on an average 6 feet high, and there were some families which had been long and closely interbred, these would be almost dwarfs, their average height during ten generations being only 4 feet 8 1/4 inches.

It should be especially observed that the average difference between the crossed and self-fertilised plants is not due to a few of the former having grown to an extraordinary height, or to a few of the self-fertilised being extremely short, but to all the crossed plants having surpassed their self-fertilised opponents, with the few following exceptions. The first occurred in the sixth generation, in which the plant named “Hero” appeared; two in the eighth generation, but the self-fertilised plants in this generation were in an anomalous condition, as they grew at first at an unusual rate and conquered for a time the opposed crossed plants; and two exceptions in the ninth generation, though one of these plants only equalled its crossed opponent. Therefore, of the seventy-three crossed plants, sixty-eight grew to a greater height than the self-fertilised plants, to which they were opposed.

In the right-hand column of figures, the difference in height between the crossed and self-fertilised plants in the successive generations is seen to fluctuate much, as might indeed have been expected from the small number of plants measured in each generation being insufficient to give a fair average. It should be remembered that the absolute height of the plants goes for nothing, as each pair was measured as soon as one of them had twined up to the summit of its rod. The great difference in the tenth generation, namely, 100 to 54, no doubt was partly accidental, though, when these plants were weighed, the difference was even greater, namely, 100 to 44. The smallest amount of difference occurred in the fourth and the eighth generations, and this was apparently due to both the crossed and self-fertilised plants having become unhealthy, which prevented the former attaining their usual degree of superiority. This was an unfortunate circumstance, but my experiments were not thus vitiated, as both lots of plants were exposed to the same conditions, whether favourable or unfavourable.

There is reason to believe that the flowers of this Ipomoea, when growing out of doors, are habitually crossed by insects, so that the first seedlings which I raised from purchased seeds were probably the offspring of a cross. I infer that this is the case, firstly from humble-bees often visiting the flowers, and from the quantity of pollen left by them on the stigmas of such flowers; and, secondly, from the plants raised from the same lot of seed varying greatly in the colour of their flowers, for as we shall hereafter see, this indicates much intercrossing. (2/3. Verlot says ‘Sur la Production des Variétés’ 1865 page 66, that certain varieties of a closely allied plant, the Convolvulus tricolor, cannot be kept pure unless grown at a distance from all other varieties.) It is, therefore, remarkable that the plants raised by me from flowers which were, in all probability, self-fertilised for the first time after many generations of crossing, should have been so markedly inferior in height to the intercrossed plants as they were, namely, as 76 to 100. As the plants which were self-fertilised in each succeeding generation necessarily became much more closely interbred in the later than in the earlier generations, it might have been expected that the difference in height between them and the crossed plants would have gone on increasing; but, so far is this from being the case, that the difference between the two sets of plants in the seventh, eighth, and ninth generations taken together is less than in the first and second generations together. When, however, we remember that the self-fertilised and crossed plants are all descended from the same mother-plant, that many of the crossed plants in each generation were related, often closely related, and that all were exposed to the same conditions, which, as we shall hereafter find, is a very important circumstance, it is not at all surprising that the difference between them should have somewhat decreased in the later generations. It is, on the contrary, an astonishing fact, that the crossed plants should have been victorious, even to a slight degree, over the self-fertilised plants of the later generations.

The much greater constitutional vigour of the crossed than of the self-fertilised plants, was proved on five occasions in various ways; namely, by exposing them, while young, to a low temperature or to a sudden change of temperature, or by growing them, under very unfavourable conditions, in competition with full-grown plants of other kinds.

With respect to the productiveness of the crossed and self-fertilised plants of the successive generations, my observations unfortunately were not made on any uniform plan, partly from the want of time, and partly from not having at first intended to observe more than a single generation. A summary of the results is here given in a tabulated form, the fertility of the crossed plants being taken as 100.

TABLE 2/18. Ratio of productiveness of crossed and self-fertilised plants. Ipomoea purpurea.

FIRST GENERATION OF CROSSED AND SELF-FERTILISED PLANTS GROWING IN COMPETITION WITH ONE ANOTHER.

Sixty-five capsules produced from flowers on five crossed plants fertilised by pollen from a distinct plant, and fifty-five capsules produced from flowers on five self-fertilised plants fertilised by their own pollen, contained seeds in the proportion of : 100 to 93.

Fifty-six spontaneously self-fertilised capsules on the above five crossed plants, and twenty-five spontaneously self-fertilised capsules on the above five self-fertilised plants, yielded seeds in the proportion of : 100 to 99.

Combining the total number of capsules produced by these plants, and the average number of seeds in each, the above crossed and self-fertilised plants yielded seeds in the proportion of : 100 to 64.

Other plants of this first generation grown under unfavourable conditions and spontaneously self-fertilised, yielded seeds in the proportion of : 100 to 45.

THIRD GENERATION OF CROSSED AND SELF-FERTILISED PLANTS.

Crossed capsules compared with self-fertilised capsules contained seeds in the ratio of : 100 to 94.

An equal number of crossed and self-fertilised plants, both spontaneously self-fertilised, produced capsules in the ratio of : 100 to 38.

And these capsules contained seeds in the ratio of : 100 to 94.

Combining these data, the productiveness of the crossed to the self-fertilised plants, both spontaneously self-fertilised, was as : 100 to 35.

FOURTH GENERATION OF CROSSED AND SELF-FERTILISED PLANTS.

Capsules from flowers on the crossed plants fertilised by pollen from another plant, and capsules from flowers on the self-fertilised plants fertilised with their own pollen, contained seeds in the proportion of : 100 to 94.

FIFTH GENERATION OF CROSSED AND SELF-FERTILISED PLANTS.

The crossed plants produced spontaneously a vast number more pods (not actually counted) than the self-fertilised, and these contained seeds in the proportion of : 100 to 89.

NINTH GENERATION OF CROSSED AND SELF-FERTILISED PLANTS.

Fourteen crossed plants, spontaneously self-fertilised, and fourteen self-fertilised plants spontaneously self-fertilised, yielded capsules (the average number of seeds per capsule not having been ascertained) in the proportion of : 100 to 26.

PLANTS DERIVED FROM A CROSSED WITH A FRESH STOCK COMPARED WITH INTERCROSSED PLANTS.

The offspring of intercrossed plants of the ninth generation, crossed by a fresh stock, compared with plants of the same stock intercrossed during ten generations, both sets of plants left uncovered and naturally fertilised, produced capsules by weight as : 100 to 51.

We see in this table that the crossed plants are always in some degree more productive than the self-fertilised plants, by whatever standard they are compared. The degree differs greatly; but this depends chiefly on whether an average was taken of the seeds alone, or of the capsules alone, or of both combined. The relative superiority of the crossed plants is chiefly due to their producing a much greater number of capsules, and not to each capsule containing a larger average number of seeds. For instance, in the third generation the crossed and self-fertilised plants produced capsules in the ratio of 100 to 38, whilst the seeds in the capsules on the crossed plants were to those on the self-fertilised plants only as 100 to 94. In the eighth generation the capsules on two self-fertilised plants (not included in table 2/18), grown in separate pots and thus not subjected to any competition, yielded the large average of 5.1 seeds. The smaller number of capsules produced by the self-fertilised plants may be in part, but not altogether, attributed to their lessened size or height; this being chiefly due to their lessened constitutional vigour, so that they were not able to compete with the crossed plants growing in the same pots. The seeds produced by the crossed flowers on the crossed plants were not always heavier than the self-fertilised seeds on the self-fertilised plants. The lighter seeds, whether produced from crossed or self-fertilised flowers, generally germinated before the heavier seeds. I may add that the crossed plants, with very few exceptions, flowered before their self-fertilised opponents, as might have been expected from their greater height and vigour.

The impaired fertility of the self-fertilised plants was shown in another way, namely, by their anthers being smaller than those in the flowers on the crossed plants. This was first observed in the seventh generation, but may have occurred earlier. Several anthers from flowers on the crossed and self-fertilised plants of the eighth generation were compared under the microscope; and those from the former were generally longer and plainly broader than the anthers of the self-fertilised plants. The quantity of pollen contained in one of the latter was, as far as could be judged by the eye, about half of that contained in one from a crossed plant. The impaired fertility of the self-fertilised plants of the eighth generation was also shown in another manner, which may often be observed in hybrids—namely, by the first-formed flowers being sterile. For instance, the fifteen first flowers on a self-fertilised plant of one of the later generations were carefully fertilised with their own pollen, and eight of them dropped off; at the same time fifteen flowers on a crossed plant growing in the same pot were self-fertilised, and only one dropped off. On two other crossed plants of the same generation, several of the earliest flowers were observed to fertilise themselves and to produce capsules. In the plants of the ninth, and I believe of some previous generations, very many of the flowers, as already stated, were slightly monstrous; and this probably was connected with their lessened fertility.

All the self-fertilised plants of the seventh generation, and I believe of one or two previous generations, produced flowers of exactly the same tint, namely, of a rich dark purple. So did all the plants, without any exception, in the three succeeding generations of self-fertilised plants; and very many were raised on account of other experiments in progress not here recorded. My attention was first called to this fact by my gardener remarking that there was no occasion to label the self-fertilised plants, as they could always be known by their colour. The flowers were as uniform in tint as those of a wild species growing in a state of nature; whether the same tint occurred, as is probable, in the earlier generations, neither my gardener nor self could recollect. The flowers on the plants which were first raised from purchased seed, as well as during the first few generations, varied much in the depth of the purple tint; many were more or less pink, and occasionally a white variety appeared. The crossed plants continued to the tenth generation to vary in the same manner as before, but to a much less degree, owing, probably, to their having become more or less closely inter-related. We must therefore attribute the extraordinary uniformity of colour in the flowers on the plants of the seventh and succeeding self-fertilised generations, to inheritance not having been interfered with by crosses during several preceding generations, in combination with the conditions of life having been very uniform.

A plant appeared in the sixth self-fertilised generation, named the Hero, which exceeded by a little in height its crossed antagonist, and which transmitted its powers of growth and increased self-fertility to its children and grandchildren. A cross between the children of Hero did not give to the grandchildren any advantage over the self-fertilised grandchildren raised from the self-fertilised children. And as far as my observations can be trusted, which were made on very unhealthy plants, the great-grandchildren raised from intercrossing the grandchildren had no advantage over the seedlings from the grandchildren the product of continued self-fertilisation; and what is far more remarkable, the great-grandchildren raised by crossing the grandchildren with a fresh stock, had no advantage over either the intercrossed or self-fertilised great-grandchildren. It thus appears that Hero and its descendants differed in constitution in an extraordinary manner from ordinary plants of the present species.

Although the plants raised during ten successive generations from crosses between distinct yet inter-related plants almost invariably exceeded in height, constitutional vigour, and fertility their self-fertilised opponents, it has been proved that seedlings raised by intercrossing flowers on the same plant are by no means superior, on the contrary are somewhat inferior in height and weight, to seedlings raised from flowers fertilised with their own pollen. This is a remarkable fact, which seems to indicate that self-fertilisation is in some manner more advantageous than crossing, unless the cross brings with it, as is generally the case, some decided and preponderant advantage; but to this subject I shall recur in a future chapter.

The benefits which so generally follow from a cross between two plants apparently depend on the two differing somewhat in constitution or character. This is shown by the seedlings from the intercrossed plants of the ninth generation, when crossed with pollen from a fresh stock, being as superior in height and almost as superior in fertility to the again intercrossed plants, as these latter were to seedlings from self-fertilised plants of the corresponding generation. We thus learn the important fact that the mere act of crossing two distinct plants, which are in some degree inter-related and which have been long subjected to nearly the same conditions, does little good as compared with that from a cross between plants belonging to different stocks or families, and which have been subjected to somewhat different conditions. We may attribute the good derived from the crossing of the intercrossed plants during the ten successive generations to their still differing somewhat in constitution or character, as was indeed proved by their flowers still differing somewhat in colour. But the several conclusions which may be deduced from the experiments on Ipomoea will be more fully considered in the final chapters, after all my other observations have been given.

CHAPTER III. SCROPHULARIACEAE, GESNERIACEAE, LABIATAE, ETC.

Mimulus luteus; height, vigour, and fertility of the crossed and
self-fertilised plants of the first four generations.
Appearance of a new, tall, and highly self-fertile variety.
Offspring from a cross between self-fertilised plants.
Effects of a cross with a fresh stock.
Effects of crossing flowers on the same plant.
Summary on Mimulus luteus.
Digitalis purpurea, superiority of the crossed plants.
Effects of crossing flowers on the same plant.
Calceolaria.
Linaria vulgaris.
Verbascum thapsus.
Vandellia nummularifolia.
Cleistogene flowers.
Gesneria pendulina.
Salvia coccinea.
Origanum vulgare, great increase of the crossed plants by stolons.
Thunbergia alata.

In the family of the Scrophulariaceae I experimented on species in the six following genera: Mimulus, Digitalis, Calceolaria, Linaria, Verbascum, and Vandellia.

[3/2. SCROPHULARIACEAE.—Mimulus luteus.

The plants which I raised from purchased seed varied greatly in the colour of their flowers, so that hardly two individuals were quite alike; the corolla being of all shades of yellow, with the most diversified blotches of purple, crimson, orange, and coppery brown. But these plants differed in no other respect. (3/1. I sent several specimens with variously coloured flowers to Kew, and Dr. Hooker informs me that they all consisted of Mimulus luteus. The flowers with much red have been named by horticulturists as var. Youngiana.) The flowers are evidently well adapted for fertilisation by the agency of insects; and in the case of a closely allied species, Mimulus rosea, I have watched bees entering the flowers, thus getting their backs well dusted with pollen; and when they entered another flower the pollen was licked off their backs by the two-lipped stigma, the lips of which are irritable and close like a forceps on the pollen-grains. If no pollen is enclosed between the lips, these open again after a time. Mr. Kitchener has ingeniously explained the use of these movements, namely, to prevent the self-fertilisation of the flower. (3/2. ‘A Year’s Botany’ 1874 page 118.) If a bee with no pollen on its back enters a flower it touches the stigma, which quickly closes, and when the bee retires dusted with pollen, it can leave none on the stigma of the same flower. But as soon as it enters any other flower, plenty of pollen is left on the stigma, which will be thus cross-fertilised. Nevertheless, if insects are excluded, the flowers fertilise themselves perfectly and produce plenty of seed; but I did not ascertain whether this is effected by the stamens increasing in length with advancing age, or by the bending down of the pistil. The chief interest in my experiments on the present species, lies in the appearance in the fourth self-fertilised generation of a variety which bore large peculiarly-coloured flowers, and grew to a greater height than the other varieties; it likewise became more highly self-fertile, so that this variety resembles the plant named Hero, which appeared in the sixth self-fertilised generation of Ipomoea.

Some flowers on one of the plants raised from the purchased seeds were fertilised with their own pollen; and others on the same plant were crossed with pollen from a distinct plant. The seeds from twelve capsules thus produced were placed in separate watch-glasses for comparison; and those from the six crossed capsules appeared to the eye hardly more numerous than those from the six self-fertilised capsules. But when the seeds were weighed, those from the crossed capsules amounted to 1.02 grain, whilst those from the self-fertilised capsules were only .81 grain; so that the former were either heavier or more numerous than the latter, in the ratio of 100 to 79.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIRST GENERATION.

Having ascertained, by leaving crossed and self-fertilised seed on damp sand, that they germinated simultaneously, both kinds were thickly sown on opposite sides of a broad and rather shallow pan; so that the two sets of seedlings, which came up at the same time, were subjected to the same unfavourable conditions. This was a bad method of treatment, but this species was one of the first on which I experimented. When the crossed seedlings were on an average half an inch high, the self-fertilised ones were only a quarter of an inch high. When grown to their full height under the above unfavourable conditions, the four tallest crossed plants averaged 7.62, and the four tallest self-fertilised 5.87 inches in height; or as 100 to 77. Ten flowers on the crossed plants were fully expanded before one on the self-fertilised plants. A few of these plants of both lots were transplanted into a large pot with plenty of good earth, and the self-fertilised plants, not now being subjected to severe competition, grew during the following year as tall as the crossed plants; but from a case which follows it is doubtful whether they would have long continued equal. Some flowers on the crossed plants were crossed with pollen from another plant, and the capsules thus produced contained a rather greater weight of seed than those on the self-fertilised plants again self-fertilised.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

Seeds from the foregoing plants, fertilised in the manner just stated, were sown on the opposite sides of a small pot (1) and came up crowded. The four tallest crossed seedlings, at the time of flowering, averaged 8 inches in height, whilst the four tallest self-fertilised plants averaged only 4 inches. Crossed seeds were sown by themselves in a second small pot, and self-fertilised seeds were sown by themselves in a third small pot so that there was no competition whatever between these two lots. Nevertheless the crossed plants grew from 1 to 2 inches higher on an average than the self-fertilised. Both lots looked equally vigorous, but the crossed plants flowered earlier and more profusely than the self-fertilised. In Pot 1, in which the two lots competed with each other, the crossed plants flowered first and produced a large number of capsules, whilst the self-fertilised produced only nineteen. The contents of twelve capsules from the crossed flowers on the crossed plants, and of twelve capsules from self-fertilised flowers on the self-fertilised plants, were placed in separate watch-glasses for comparison; and the crossed seeds seemed more numerous by half than the self-fertilised.

The plants on both sides of Pot 1, after they had seeded, were cut down and transplanted into a large pot with plenty of good earth, and on the following spring, when they had grown to a height of between 5 and 6 inches, the two lots were equal, as occurred in a similar experiment in the last generation. But after some weeks the crossed plants exceeded the self-fertilised ones on the opposite side of the same pot, though not nearly to so great a degree as before, when they were subjected to very severe competition.

CROSSED AND SELF-FERTILISED PLANTS OF THE THIRD GENERATION.

Crossed seeds from the crossed plants, and self-fertilised seeds from the self-fertilised plants of the last generation, were sown thickly on opposite sides of a small pot, Number 1. The two tallest plants on each side were measured after they had flowered, and the two crossed ones were 12 and 7 1/2 inches, and the two self-fertilised ones 8 and 5 1/2 inches in height; that is, in the ratio of 100 to 69. Twenty flowers on the crossed plants were again crossed and produced twenty capsules; ten of which contained 1.33 grain weight of seeds. Thirty flowers on the self-fertilised plants were again self-fertilised and produced twenty-six capsules; ten of the best of which (many being very poor) contained only .87 grain weight of seeds; that is, in the ratio of 100 to 65 by weight.

The superiority of the crossed over the self-fertilised plants was proved in various ways. Self-fertilised seeds were sown on one side of a pot, and two days afterwards crossed seeds on the opposite side. The two lots of seedlings were equal until they were above half an inch high; but when fully grown the two tallest crossed plants attained a height of 12 1/2 and 8 3/4 inches, whilst the two tallest self-fertilised plants were only 8 and 5 1/2 inches high.

In a third pot, crossed seeds were sown four days after the self-fertilised, and the seedlings from the latter had at first, as might have been expected, an advantage; but when the two lots were between 5 and 6 inches in height, they were equal, and ultimately the three tallest crossed plants were 11, 10, and 8 inches, whilst the three tallest self-fertilised were 12, 8 1/2, and 7 1/2 inches in height. So that there was not much difference between them, the crossed plants having an average advantage of only the third of an inch. The plants were cut down, and without being disturbed were transplanted into a larger pot. Thus the two lots started fair on the following spring, and now the crossed plants showed their inherent superiority, for the two tallest were 13 inches, whilst the two tallest self-fertilised plants were only 11 and 8 1/2 inches in height; or as 100 to 75. The two lots were allowed to fertilise themselves spontaneously: the crossed plants produced a large number of capsules, whilst the self-fertilised produced very few and poor ones. The seeds from eight of the capsules on the crossed plants weighed .65 grain, whilst those from eight of the capsules on the self-fertilised plants weighed only .22 grain; or as 100 to 34.

The crossed plants in the above three pots, as in almost all the previous experiments, flowered before the self-fertilised. This occurred even in the third pot in which the crossed seeds were sown four days after the self-fertilised seeds.

Lastly, seeds of both lots were sown on opposite sides of a large pot in which a Fuchsia had long been growing, so that the earth was full of roots. Both lots grew miserably; but the crossed seedlings had an advantage at all times, and ultimately attained to a height of 3 1/2 inches, whilst the self-fertilised seedlings never exceeded 1 inch. The several foregoing experiments prove in a decisive manner the superiority in constitutional vigour of the crossed over the self-fertilised plants.

In the three generations now described and taken together, the average height of the ten tallest crossed plants was 8.19 inches, and that of the ten tallest self-fertilised plants 5.29 inches (the plants having been grown in small pots), or as 100 to 65.

In the next or fourth self-fertilised generation, several plants of a new and tall variety appeared, which increased in the later self-fertilised generations, owing to its great self-fertility, to the complete exclusion of the original kinds. The same variety also appeared amongst the crossed plants, but as it was not at first regarded with any particular attention, I know not how far it was used for raising the intercrossed plants; and in the later crossed generations it was rarely present. Owing to the appearance of this tall variety, the comparison of the crossed and self-fertilised plants of the fifth and succeeding generations was rendered unfair, as all the self-fertilised and only a few or none of the crossed plants consisted of it. Nevertheless, the results of the later experiments are in some respects well worth giving.

CROSSED AND SELF-FERTILISED PLANTS OF THE FOURTH GENERATION.

Seeds of the two kinds, produced in the usual way from the two sets of plants of the third generation, were sown on opposite sides of two pots (1 and 2); but the seedlings were not thinned enough and did not grow well. Many of the self-fertilised plants, especially in one of the pots, consisted of the new and tall variety above referred to, which bore large and almost white flowers marked with crimson blotches. I will call it the WHITE VARIETY. I believe that it first appeared amongst both the crossed and self-fertilised plants of the last generation; but neither my gardener nor myself could remember any such variety in the seedlings raised from the purchased seed. It must therefore have arisen either through ordinary variation, or, judging from its appearance amongst both the crossed and self-fertilised plants, more probably through reversion to a formerly existing variety.

In Pot 1 the tallest crossed plant was 8 1/2 inches, and the tallest self-fertilised 5 inches in height. In Pot 2, the tallest crossed plant was 6 1/2 inches, and the tallest self-fertilised plant, which consisted of the white variety, 7 inches in height; and this was the first instance in my experiments on Mimulus in which the tallest self-fertilised plant exceeded the tallest crossed. Nevertheless, the two tallest crossed plants taken together were to the two tallest self-fertilised plants in height as 100 to 80. As yet the crossed plants were superior to the self-fertilised in fertility; for twelve flowers on the crossed plants were crossed and yielded ten capsules, the seeds of which weighed 1.71 grain. Twenty flowers on the self-fertilised plants were self-fertilised, and produced fifteen capsules, all appearing poor; and the seeds from ten of them weighed only .68 grain, so that from an equal number of capsules the crossed seeds were to the self-fertilised in weight as 100 to 40.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIFTH GENERATION.

Seeds from both lots of the fourth generation, fertilised in the usual manner, were sown on opposite sides of three pots. When the seedlings flowered, most of the self-fertilised plants were found to consist of the tall white variety. Several of the crossed plants in Pot 1 likewise belonged to this variety, as did a very few in Pots 2 and 3. The tallest crossed plant in Pot 1 was 7 inches, and the tallest self-fertilised plant on the opposite side 8 inches; in Pots 2 and 3 the tallest crossed were 4 1/2 and 5 1/2, and the tallest self-fertilised 7 and 6 1/2 inches in height; so that the average height of the tallest plants in the two lots was as 100 for the crossed to 126 for the self-fertilised; and thus we have a complete reversal of what occurred in the four previous generations. Nevertheless, in all three pots the crossed plants retained their habit of flowering before the self-fertilised. The plants were unhealthy from being crowded and from the extreme heat of the season, and were in consequence more or less sterile; but the crossed plants were somewhat less sterile than the self-fertilised plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SIXTH GENERATION.

Seeds from plants of the fifth generation crossed and self-fertilised in the usual manner were sown on opposite sides of several pots. On the self-fertilised side every single plant belonged to the tall white variety. On the crossed side some plants belonged to this variety, but the greater number approached in character to the old and shorter kinds with smaller yellowish flowers blotched with coppery brown. When the plants on both sides were from 2 to 3 inches in height they were equal, but when fully grown the self-fertilised were decidedly the tallest and finest plants, but, from want of time, they were not actually measured. In half the pots the first plant which flowered was a self-fertilised one, and in the other half a crossed one. And now another remarkable change was clearly perceived, namely, that the self-fertilised plants had become more self-fertile than the crossed. The pots were all put under a net to exclude insects, and the crossed plants produced spontaneously only fifty-five capsules, whilst the self-fertilised plants produced eighty-one capsules, or as 100 to 147. The seeds from nine capsules of both lots were placed in separate watch-glasses for comparison, and the self-fertilised appeared rather the more numerous. Besides these spontaneously self-fertilised capsules, twenty flowers on the crossed plants again crossed yielded sixteen capsules; twenty-five flowers on the self-fertilised plants again self-fertilised yielded seventeen capsules, and this is a larger proportional number of capsules than was produced by the self-fertilised flowers on the self-fertilised plants in the previous generations. The contents of ten capsules of both these lots were compared in separate watch-glasses, and the seeds from the self-fertilised appeared decidedly more numerous than those from the crossed plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SEVENTH GENERATION.

Crossed and self-fertilised seeds from the crossed and self-fertilised plants of the sixth generation were sown in the usual manner on opposite sides of three pots, and the seedlings were well and equally thinned. Every one of the self-fertilised plants (and many were raised) in this, as well as in the eighth and ninth generations, belonged to the tall white variety. Their uniformity of character, in comparison with the seedlings first raised from the purchased seed, was quite remarkable. On the other hand, the crossed plants differed much in the tints of their flowers, but not, I think, to so great a degree as those first raised. I determined this time to measure the plants on both sides carefully. The self-fertilised seedlings came up rather before the crossed, but both lots were for a time of equal height. When first measured, the average height of the six tallest crossed plants in the three pots was 7.02, and that of the six tallest self-fertilised plants 8.97 inches, or as 100 to 128. When fully grown the same plants were again measured, with the result shown in Table 3/18.

TABLE 3/18. Mimulus luteus (Seventh Generation).

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 11 2/8 : 19 1/8. Pot 1 : 11 7/8 : 18.

Pot 2 : 12 6/8 : 18 2/8. Pot 2 : 11 2/8 : 14 6/8.

Pot 3 : 9 6/8 : 12 6/8. Pot 3 : 11 6/8 : 11.

Total : 68.63 : 93.88.

The average height of the six crossed is here 11.43, and that of the six self-fertilised 15.64, or as 100 to 137.

As it is now evident that the tall white variety transmitted its characters faithfully, and as the self-fertilised plants consisted exclusively of this variety, it was manifest that they would always exceed in height the crossed plants which belonged chiefly to the original shorter varieties. This line of experiment was therefore discontinued, and I tried whether intercrossing two self-fertilised plants of the sixth generation, growing in distinct pots, would give their offspring any advantage over the offspring of flowers on one of the same plants fertilised with their own pollen. These latter seedlings formed the seventh generation of self-fertilised plants, like those in the right hand column in Table 3/18; the crossed plants were the product of six previous self-fertilised generations with an intercross in the last generation. The seeds were allowed to germinate on sand, and were planted in pairs on opposite sides of four pots, all the remaining seeds being sown crowded on opposite sides of Pot 5 in Table 3/19; the three tallest on each side in this latter pot being alone measured. All the plants were twice measured—the first time whilst young, and the average height of the crossed plants to that of the self-fertilised was then as 100 to 122. When fully grown they were again measured, as in Table 3/19.

TABLE 3/19. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Intercrossed Plants from Self-fertilised Plants of the Sixth Generation.

Column 3: Self-fertilised Plants of the Seventh Generation.

Pot 1 : 12 6/8 : 15 2/8. Pot 1 : 10 4/8 : 11 5/8. Pot 1 : 10 : 11. Pot 1 : 14 5/8 : 11.

Pot 2 : 10 2/8 : 11 3/8. Pot 2 : 7 6/8 : 11 4/8. Pot 2 : 12 1/8 : 8 5/8. Pot 2 : 7 : 14 3/8.

Pot 3 : 13 5/8 : 10 3/8. Pot 3 : 12 2/8 : 11 6/8.

Pot 4 : 7 1/8 : 14 6/8. Pot 4 : 8 2/8 : 7. Pot 4 : 7 2/8 : 8.

Pot 5 : 8 5/8 : 10 2/8 Pot 5 : 9 : 9 3/8. Pot 5 : 8 2/8 : 9 2/8. Crowded.

Total : 159.38 : 175.50.

The average height of the sixteen intercrossed plants is here 9.96 inches, and that of the sixteen self-fertilised plants 10.96, or as 100 to 110; so that the intercrossed plants, the progenitors of which had been self-fertilised for the six previous generations, and had been exposed during the whole time to remarkably uniform conditions, were somewhat inferior in height to the plants of the seventh self-fertilised generation. But as we shall presently see that a similar experiment made after two additional generations of self-fertilisation gave a different result, I know not how far to trust the present one. In three of the five pots in Table 3/19 a self-fertilised plant flowered first, and in the other two a crossed plant. These self-fertilised plants were remarkably fertile, for twenty flowers fertilised with their own pollen produced no less than nineteen very fine capsules!

THE EFFECTS OF A CROSS WITH A DISTINCT STOCK.

Some flowers on the self-fertilised plants in Pot 4 in Table 3/19 were fertilised with their own pollen, and plants of the eighth self-fertilised generation were thus raised, merely to serve as parents in the following experiment. Several flowers on these plants were allowed to fertilise themselves spontaneously (insects being of course excluded), and the plants raised from these seeds formed the ninth self-fertilised generation; they consisted wholly of the tall white variety with crimson blotches. Other flowers on the same plants of the eighth self-fertilised generation were crossed with pollen taken from another plant of the same lot; so that the seedlings thus raised were the offspring of eight previous generations of self-fertilisation with an intercross in the last generation; these I will call the INTERCROSSED PLANTS. Lastly, other flowers on the same plants of the eighth self-fertilised generation were crossed with pollen taken from plants which had been raised from seed procured from a garden at Chelsea. The Chelsea plants bore yellow flowers blotched with red, but differed in no other respect. They had been grown out of doors, whilst mine had been cultivated in pots in the greenhouse for the last eight generations, and in a different kind of soil. The seedlings raised from this cross with a wholly different stock may be called the CHELSEA-CROSSED. The three lots of seeds thus obtained were allowed to germinate on bare sand; and whenever a seed in all three lots, or in only two, germinated at the same time, they were planted in pots superficially divided into three or two compartments. The remaining seeds, whether or not in a state of germination, were thickly sown in three divisions in a large pot, 10, in Table 3/20. When the plants had grown to their full height they were measured, as shown in Table 3/20; but only the three tallest plants in each of the three divisions in Pot 10 were measured.

TABLE 3/20. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants from Self-fertilised Plants of the Eighth Generation crossed by Chelsea Plants.

Column 3: Plants from an intercross between the Plants of the Eighth Self-fertilised Generation.

Column 4: Self-fertilised Plants of the Ninth Generation from Plants of the Eighth Self-fertilised Generation.

Pot 1 : 30 7/8 : 14 : 9 4/8. Pot 1 : 28 3/8 : 13 6/8 : 10 5/8. Pot 1 : — : 13 7/8 : 10.

Pot 2 : 20 6/8 : 11 4/8 : 11 6/8. Pot 2 : 22 2/8 : 12 : 12 3/8. Pot 2 : — : 9 1/8 : —.

Pot 3 : 23 6/8 : 12 2/8 : 8 5/8. Pot 3 : 24 1/8 : — : 11 4/8. Pot 3 : 25 6/8 : — : 6 7/8.

Pot 4 : 22 5/8 : 9 2/8 : 4. Pot 4 : 22 : 8 1/8 : 13 3/8. Pot 4 : 17 : — : 11.

Pot 5 : 22 3/8 : 9 : 4 4/8. Pot 5 : 19 5/8 : 11 : 13. Pot 5 : 23 4/8 : — : 13 4/8.

Pot 6 : 28 2/8 : 18 6/8 : 12. Pot 6 : 22 : 7 : 16 1/8. Pot 6 : — : 12 4/8 : —.

Pot 7 : 12 4/8 : 15 : —. Pot 7 : 24 3/8 : 12 3/8 : —. Pot 7 : 20 4/8 : 11 2/8 : —. Pot 7 : 26 4/8 : 15 2/8 : —.

Pot 8 : 17 2/8 : 13 3/8 : —. Pot 8 : 22 6/8 : 14 5/8 : —. Pot 8 : 27 : 14 3/8 : —.

Pot 9 : 22 6/8 : 11 6/8 : —. Pot 9 : 6 : 17 : —. Pot 9 : 20 2/8 : 14 7/8 : —.

Pot 10 : 18 1/8 : 9 2/8 : 10 3/8. Pot 10 : 16 5/8 : 8 2/8 : 8 1/8. Pot 10 : 17 4/8 : 10 : 11 2/8. Crowded plants.

Total : 605.38 : 329.50 : 198.50.

In this table the average height of the twenty-eight Chelsea-crossed plants is 21.62 inches; that of the twenty-seven intercrossed plants 12.2; and that of the nineteen self-fertilised 10.44. But with respect to the latter it will be the fairest plan to strike out two dwarfed ones (only 4 inches in height), so as not to exaggerate the inferiority of the self-fertilised plants; and this will raise the average height of the seventeen remaining self-fertilised plants to 11.2 inches. Therefore the Chelsea-crossed are to the intercrossed in height as 100 to 56; the Chelsea-crossed to the self-fertilised as 100 to 52; and the intercrossed to the self-fertilised as 100 to 92. We thus see how immensely superior in height the Chelsea-crossed are to the intercrossed and to the self-fertilised plants. They began to show their superiority when only one inch high. They were also, when fully grown, much more branched with larger leaves and somewhat larger flowers than the plants of the other two lots, so that if they had been weighed, the ratio would certainly have been much higher than that of 100 to 56 and 52.

The intercrossed plants are here to the self-fertilised in height as 100 to 92; whereas in the analogous experiment given in Table 3/19 the intercrossed plants from the self-fertilised plants of the sixth generation were inferior in height to the self-fertilised plants in the ratio of 100 to 110. I doubt whether this discordance in the results of the two experiments can be explained by the self-fertilised plants in the present case having been raised from spontaneously self-fertilised seeds, whereas in the former case they were raised from artificially self-fertilised seeds; nor by the present plants having been self-fertilised during two additional generations, though this is a more probable explanation.

With respect to fertility, the twenty-eight Chelsea-crossed plants produced 272 capsules; the twenty-seven intercrossed plants produced 24; and the seventeen self-fertilised plants 17 capsules. All the plants were left uncovered so as to be naturally fertilised, and empty capsules were rejected.

Therefore 20 Chelsea-crossed plants would have produced 194.29 capsules.

Therefore 20 Intercrossed plants would have produced 17.77 capsules.

Therefore 20 Self-fertilised plants would have produced 20.00 capsules.

The seeds contained in 8 capsules from the Chelsea-crossed plants weighed 1.1 grains.

The seeds contained in 8 capsules from the Intercrossed plants weighed 0.51 grains.

The seeds contained in 8 capsules from the Self-fertilised plants weighed 0.33 grains.

If we combine the number of capsules produced together with the average weight of contained seeds, we get the following extraordinary ratios:

Weight of seed produced by the same number of Chelsea-crossed and intercrossed plants as 100 to 4.

Weight of seed produced by the same number of Chelsea-crossed and self-fertilised plants as 100 to 3.

Weight of seeds produced by the same number of intercrossed and self-fertilised plants as 100 to 73.

It is also a remarkable fact that the Chelsea-crossed plants exceeded the two other lots in hardiness, as greatly as they did in height, luxuriance, and fertility. In the early autumn most of the pots were bedded out in the open ground; and this always injures plants which have been long kept in a warm greenhouse. All three lots consequently suffered greatly, but the Chelsea-crossed plants much less than the other two lots. On the 3rd of October the Chelsea-crossed plants began to flower again, and continued to do so for some time; whilst not a single flower was produced by the plants of the other two lots, the stems of which were cut almost down to the ground and seemed half dead. Early in December there was a sharp frost, and the stems of Chelsea-crossed were now cut down; but on the 23rd of December they began to shoot up again from the roots, whilst all the plants of the other two lots were quite dead.

Although several of the self-fertilised seeds, from which the plants in the right hand column in Table 3/20 were raised, germinated (and were of course rejected) before any of those of the other two lots, yet in only one of the ten pots did a self-fertilised plant flower before the Chelsea-crossed or the intercrossed plants growing in the same pots. The plants of these two latter lots flowered at the same time, though the Chelsea-crossed grew so much taller and more vigorously than the intercrossed.

As already stated, the flowers of the plants originally raised from the Chelsea seeds were yellow; and it deserves notice that every one of the twenty-eight seedlings raised from the tall white variety fertilised, without being castrated, with pollen from the Chelsea plants, produced yellow flowers; and this shows how prepotent this colour, which is the natural one of the species, is over the white colour.

THE EFFECTS ON THE OFFSPRING OF INTERCROSSING FLOWERS ON THE SAME PLANT, INSTEAD OF CROSSING DISTINCT INDIVIDUALS.

In all the foregoing experiments the crossed plants were the product of a cross between distinct plants. I now selected a very vigorous plant in Table 3/20, raised by fertilising a plant of the eighth self-fertilised generation with pollen from the Chelsea stock. Several flowers on this plant were crossed with pollen from other flowers on the same plant, and several other flowers were fertilised with their own pollen. The seed thus produced was allowed to germinate on bare sand; and the seedlings were planted in the usual manner on the opposite sides of six pots. All the remaining seeds, whether or not in a state of germination, were sown thickly in Pot 7; the three tallest plants on each side of this latter pot being alone measured. As I was in a hurry to learn the result, some of these seeds were sown late in the autumn, but the plants grew so irregularly during the winter, that one crossed plant was 28 1/2 inches, and two others only 4, or less than 4 inches in height, as may be seen in Table 3/21. Under such circumstances, as I have observed in many other cases, the result is not in the least trustworthy; nevertheless I feel bound to give the measurements.

TABLE 3/21. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 17 : 17. Pot 1 : 9 : 3 1/8.

Pot 2 : 28 2/8 : 19 1/8. Pot 2 : 16 4/8 : 6. Pot 2 : 13 5/8 : 2.

Pot 3 : 4 : 15 6/8. Pot 3 : 2 2/8 : 10.

Pot 4 : 23 4/8 : 6 2/8. Pot 4 : 15 4/8 : 7 1/8.

Pot 5 : 7 : 13 4/8.

Pot 6 : 18 3/8 : 1 4/8. Pot 6 : 11 : 2.

Pot 7 : 21 : 15 1/8. Pot 7 : 11 6/8 : 11. Pot 7 : 12 1/8 : 11 2/8. Crowded.

Total : 210.88 : 140.75.

The fifteen crossed plants here average 14.05, and the fifteen self-fertilised plants 9.38 in height, or as 100 to 67. But if all the plants under ten inches in height are struck out, the ratio of the eleven crossed plants to the eight self-fertilised plants is as 100 to 82.

On the following spring, some remaining seeds of the two lots were treated in exactly the same manner; and the measurements of the seedlings are given in Table 3/22.

TABLE 3/22. Mimulus luteus.

Heights of Plants in inches:

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 15 1/8 : 19 1/8. Pot 1 : 12 : 20 5/8. Pot 1 : 10 1/8 : 12 6/8.

Pot 2 : 16 2/8 : 11 2/8. Pot 2 : 13 5/8 : 19 3/8. Pot 2 : 20 1/8 : 17 4/8.

Pot 3 : 18 7/8 : 12 6/8. Pot 3 : 15 : 15 6/8. Pot 3 : 13 7/8 : 17.

Pot 4 : 19 2/8 : 16 2/8. Pot 4 : 19 6/8 : 21 5/8.

Pot 5 : 25 3/8 : 22 5/8.

Pot 6 : 15 : 19 5/8. Pot 6 : 20 2/8 : 16 2/8. Pot 6 : 27 2/8 : 19 5/8.

Pot 7 : 7 6/8 : 7 6/8. Pot 7 : 14 : 8. Pot 7 : 13 4/8 : 7.

Pot 8 : 18 2/8 : 20 3/8. Pot 8 : 18 6/8 : 17 6/8. Pot 8 : 18 3/8 : 15 4/8. Pot 8 : 18 3/8 : 15 1/8. Crowded.

Total : 370.88 : 353.63.

Here the average height of the twenty-two crossed plants is 16.85, and that of the twenty-two self-fertilised plants 16.07; or as 100 to 95. But if four of the plants in Pot 7, which are much shorter than any of the others, are struck out (and this would be the fairest plan), the twenty-one crossed are to the nineteen self-fertilised plants in height as 100 to 100.6—that is, are equal. All the plants, except the crowded ones in Pot 8, after being measured were cut down, and the eighteen crossed plants weighed 10 ounces, whilst the same number of self-fertilised plants weighed 10 1/4 ounces, or as 100 to 102.5; but if the dwarfed plants in Pot 7 had been excluded, the self-fertilised would have exceeded the crossed in weight in a higher ratio. In all the previous experiments in which seedlings were raised from a cross between distinct plants, and were put into competition with self-fertilised plants, the former generally flowered first; but in the present case, in seven out of the eight pots a self-fertilised plant flowered before a crossed one on the opposite side. Considering all the evidence with respect to the plants in Table3/ 22, a cross between two flowers on the same plant seems to give no advantage to the offspring thus produced, the self-fertilised plants being in weight superior. But this conclusion cannot be absolutely trusted, owing to the measurements given in Table 3/21, though these latter, from the cause already assigned, are very much less trustworthy than the present ones.]

A SUMMARY OF OBSERVATIONS ON Mimulus luteus.

In the three first generations of crossed and self-fertilised plants, the tallest plants alone on each side of the several pots were measured; and the average height of the ten crossed to that of the ten self-fertilised plants was as 100 to 64. The crossed were also much more fertile than the self-fertilised, and so much more vigorous that they exceeded them in height, even when sown on the opposite side of the same pot after an interval of four days. The same superiority was likewise shown in a remarkable manner when both kinds of seeds were sown on the opposite sides of a pot with very poor earth full of the roots of another plant. In one instance crossed and self-fertilised seedlings, grown in rich soil and not put into competition with each other, attained to an equal height. When we come to the fourth generation the two tallest crossed plants taken together exceeded by only a little the two tallest self-fertilised plants, and one of the latter beat its crossed opponent,—a circumstance which had not occurred in the previous generations. This victorious self-fertilised plant consisted of a new white-flowered variety, which grew taller than the old yellowish varieties. From the first it seemed to be rather more fertile, when self-fertilised, than the old varieties, and in the succeeding self-fertilised generations became more and more self-fertile. In the sixth generation the self-fertilised plants of this variety compared with the crossed plants produced capsules in the proportion of 147 to 100, both lots being allowed to fertilise themselves spontaneously. In the seventh generation twenty flowers on one of these plants artificially self-fertilised yielded no less than nineteen very fine capsules!

This variety transmitted its characters so faithfully to all the succeeding self-fertilised generations, up to the last or ninth, that all the many plants which were raised presented a complete uniformity of character; thus offering a remarkable contrast with the seedlings raised from the purchased seeds. Yet this variety retained to the last a latent tendency to produce yellow flowers; for when a plant of the eighth self-fertilised generation was crossed with pollen from a yellow-flowered plant of the Chelsea stock, every single seedling bore yellow flowers. A similar variety, at least in the colour of its flowers, also appeared amongst the crossed plants of the third generation. No attention was at first paid to it, and I know not how far it was at first used either for crossing or self-fertilisation. In the fifth generation most of the self-fertilised plants, and in the sixth and all the succeeding generations every single plant consisted of this variety; and this no doubt was partly due to its great and increasing self-fertility. On the other hand, it disappeared from amongst the crossed plants in the later generations; and this was probably due to the continued intercrossing of the several plants. From the tallness of this variety, the self-fertilised plants exceeded the crossed plants in height in all the generations from the fifth to the seventh inclusive; and no doubt would have done so in the later generations, had they been grown in competition with one another. In the fifth generation the crossed plants were in height to the self-fertilised, as 100 to 126; in the sixth, as 100 to 147; and in the seventh generation, as 100 to 137. This excess of height may be attributed not only to this variety naturally growing taller than the other plants, but to its possessing a peculiar constitution, so that it did not suffer from continued self-fertilisation.

This variety presents a strikingly analogous case to that of the plant called the Hero, which appeared in the sixth self-fertilised generation of Ipomoea. If the seeds produced by Hero had been as greatly in excess of those produced by the other plants, as was the case with Mimulus, and if all the seeds had been mingled together, the offspring of Hero would have increased to the entire exclusion of the ordinary plants in the later self-fertilised generations, and from naturally growing taller would have exceeded the crossed plants in height in each succeeding generation.

Some of the self-fertilised plants of the sixth generation were intercrossed, as were some in the eighth generation; and the seedlings from these crosses were grown in competition with self-fertilised plants of the two corresponding generations. In the first trial the intercrossed plants were less fertile than the self-fertilised, and less tall in the ratio of 100 to 110. In the second trial, the intercrossed plants were more fertile than the self-fertilised in the ratio of 100 to 73, and taller in the ratio of 100 to 92. Notwithstanding that the self-fertilised plants in the second trial were the product of two additional generations of self-fertilisation, I cannot understand this discordance in the results of the two analogous experiments.

The most important of all the experiments on Mimulus are those in which flowers on plants of the eighth self-fertilised generation were again self-fertilised; other flowers on distinct plants of the same lot were intercrossed; and others were crossed with a new stock of plants from Chelsea. The Chelsea-crossed seedlings were to the intercrossed in height as 100 to 56, and in fertility as 100 to 4; and they were to the self-fertilised plants, in height as 100 to 52, and in fertility as 100 to 3. These Chelsea-crossed plants were also much more hardy than the plants of the other two lots; so that altogether the gain from the cross with a fresh stock was wonderfully great.

Lastly, seedlings raised from a cross between flowers on the same plant were not superior to those from flowers fertilised with their own pollen; but this result cannot be absolutely trusted, owing to some previous observations, which, however, were made under very unfavourable circumstances.

[Digitalis purpurea.

The flowers of the common Foxglove are proterandrous; that is, the pollen is mature and mostly shed before the stigma of the same flower is ready for fertilisation. This is effected by the larger humble-bees, which, whilst in search of nectar, carry pollen from flower to flower. The two upper and longer stamens shed their pollen before the two lower and shorter ones. The meaning of this fact probably is, as Dr. Ogle remarks, that the anthers of the longer stamens stand near to the stigma, so that they would be the most likely to fertilise it (3/3. ‘Popular Science Review’ January 1870 page 50.); and as it is an advantage to avoid self-fertilisation, they shed their pollen first, thus lessening the chance. There is, however, but little danger of self-fertilisation until the bifid stigma opens; for Hildebrand found that pollen placed on the stigma before it had opened produced no effect. (3/4. ‘Geschlechter-Vertheilung bei den Pflanzen’ 1867 page 20.) The anthers, which are large, stand at first transversely with respect to the tubular corolla, and if they were to dehisce in this position they would, as Dr. Ogle also remarks, smear with pollen the whole back and sides of an entering humble-bee in a useless manner; but the anthers twist round and place themselves longitudinally before they dehisce. The lower and inner side of the mouth of the corolla is thickly clothed with hairs, and these collect so much of the fallen pollen that I have seen the under surface of a humble-bee thickly dusted with it; but this can never be applied to the stigma, as the bees in retreating do not turn their under surfaces upwards. I was therefore puzzled whether these hairs were of any use; but Mr. Belt has, I think, explained their use: the smaller kinds of bees are not fitted to fertilise the flowers, and if they were allowed to enter easily they would steal much nectar, and fewer large bees would haunt the flowers. Humble-bees can crawl into the dependent flowers with the greatest ease, using the “hairs as footholds while sucking the honey; but the smaller bees are impeded by them, and when, having at length struggled through them, they reach the slippery precipice above, they are completely baffled.” Mr. Belt says that he watched many flowers during a whole season in North Wales, and “only once saw a small bee reach the nectary, though many were seen trying in vain to do so.” (3/5. ‘The Naturalist in Nicaragua’ 1874 page 132. But it appears from H. Muller ‘Die Befruchtung der Blumen’ 1873 page 285, that small insects sometimes succeed in entering the flowers.)

I covered a plant growing in its native soil in North Wales with a net, and fertilised six flowers each with its own pollen, and six others with pollen from a distinct plant growing within the distance of a few feet. The covered plant was occasionally shaken with violence, so as to imitate the effects of a gale of wind, and thus to facilitate as far as possible self-fertilisation. It bore ninety-two flowers (besides the dozen artificially fertilised), and of these only twenty-four produced capsules; whereas almost all the flowers on the surrounding uncovered plants were fruitful. Of the twenty-four spontaneously self-fertilised capsules, only two contained their full complement of seed; six contained a moderate supply; and the remaining sixteen extremely few seeds. A little pollen adhering to the anthers after they had dehisced, and accidentally falling on the stigma when mature, must have been the means by which the above twenty-four flowers were partially self-fertilised; for the margins of the corolla in withering do not curl inwards, nor do the flowers in dropping off turn round on their axes, so as to bring the pollen-covered hairs, with which the lower surface is clothed, into contact with the stigma—by either of which means self-fertilisation might be effected.

Seeds from the above crossed and self-fertilised capsules, after germinating on bare sand, were planted in pairs on the opposite sides of five moderately-sized pots, which were kept in the greenhouse. The plants after a time appeared starved, and were therefore, without being disturbed, turned out of their pots, and planted in the open ground in two close parallel rows. They were thus subjected to tolerably severe competition with one another; but not nearly so severe as if they had been left in the pots. At the time when they were turned out, their leaves were between 5 and 8 inches in length, and the longest leaf on the finest plant on each side of each pot was measured, with the result that the leaves of the crossed plants exceeded, on an average, those of the self-fertilised plants by .4 of an inch.

In the following summer the tallest flower-stem on each plant, when fully grown, was measured. There were seventeen crossed plants; but one did not produce a flower-stem. There were also, originally, seventeen self-fertilised plants, but these had such poor constitutions that no less than nine died in the course of the winter and spring, leaving only eight to be measured, as in Table 3/23.

TABLE 3/23. Digitalis purpurea.

The tallest Flower-stem on each Plant measured in inches: 0 means that the Plant died before a Flower-stem was produced.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 53 6/8 : 27 4/8. Pot 1 : 57 4/8 : 55 6/8. Pot 1 : 57 6/8 : 0. Pot 1 : 65 : 0.

Pot 2 : 34 4/8 : 39. Pot 2 : 52 4/8 : 32. Pot 2 : 63 6/8 : 21.

Pot 3 : 57 4/8 : 53 4/8. Pot 3 : 53 4/8 : 0. Pot 3 : 50 6/8 : 0. Pot 3 : 37 2/8 : 0.

Pot 4 : 64 4/8 : 34 4/8. Pot 4 : 37 4/8 : 23 6/8. Pot 4 : — : 0.

Pot 5 : 53 : 0. Pot 5 : 47 6/8 : 0. Pot 5 : 34 6/8 : 0.

Total : 821.25 : 287.00.

The average height of the flower-stems of the sixteen crossed plants is here 51.33 inches; and that of the eight self-fertilised plants, 35.87; or as 100 to 70. But this difference in height does not give at all a fair idea of the vast superiority of the crossed plants. These latter produced altogether sixty-four flower-stems, each plant producing, on an average, exactly four flower-stems, whereas the eight self-fertilised plants produced only fifteen flower-stems, each producing an average only of 1.87 stems, and these had a less luxuriant appearance. We may put the result in another way: the number of flower-stems on the crossed plants was to those on an equal number of self-fertilised plants as 100 to 48.

Three crossed seeds in a state of germination were also planted in three separate pots; and three self-fertilised seeds in the same state in three other pots. These plants were therefore at first exposed to no competition with one another, and when turned out of their pots into the open ground they were planted at a moderate distance apart, so that they were exposed to much less severe competition than in the last case. The longest leaves on the three crossed plants, when turned out, exceeded those on the self-fertilised plants by a mere trifle, namely, on an average by .17 of an inch. When fully grown the three crossed plants produced twenty-six flower-stems; the two tallest of which on each plant were on an average 54.04 inches in height. The three self-fertilised plants produced twenty-three flower-stems, the two tallest of which on each plant had an average height of 46.18 inches. So that the difference between these two lots, which hardly competed together, is much less than in the last case when there was moderately severe competition, namely, as 100 to 85, instead of as 100 to 70.

THE EFFECTS ON THE OFFSPRING OF INTERCROSSING DIFFERENT FLOWERS ON THE SAME PLANT, INSTEAD OF CROSSING DISTINCT INDIVIDUALS.

A fine plant growing in my garden (one of the foregoing seedlings) was covered with a net, and six flowers were crossed with pollen from another flower on the same plant, and six others were fertilised with their own pollen. All produced good capsules. The seeds from each were placed in separate watch-glasses, and no difference could be perceived by the eye between the two lots of seeds; and when they were weighed there was no difference of any significance, as the seeds from the self-fertilised capsules weighed 7.65 grains, whilst those from the crossed capsules weighed 7.7 grains. Therefore the sterility of the present species, when insects are excluded, is not due to the impotence of pollen on the stigma of the same flower. Both lots of seeds and seedlings were treated in exactly the same manner as in Table 3/23, excepting that after the pairs of germinating seeds had been planted on the opposite sides of eight pots, all the remaining seeds were thickly sown on the opposite sides of Pots 9 and 10 in Table 3/24. The young plants during the following spring were turned out of their pots, without being disturbed, and planted in the open ground in two rows, not very close together, so that they were subjected to only moderately severe competition with one another. Very differently to what occurred in the first experiment, when the plants were subjected to somewhat severe mutual competition, an equal number on each side either died or did not produce flower-stems. The tallest flower-stems on the surviving plants were measured, as shown in Table 3/24.

TABLE 3/24. Digitalis purpurea.

The tallest Flower-stem on each Plant measured in inches: 0 signifies that the Plant died, or did not produce a Flower-stem.

Column 1: Number (Name) of Pot.

Column 2: Plants raised from a Cross between different Flowers on the same Plant.

Column 3: Plants raised from Flowers fertilised with their own Pollen.

Pot 1 : 49 4/8 : 45 5/8. Pot 1 : 46 7/8 : 52. Pot 1 : 43 6/8 : 0.

Pot 2 : 38 4/8 : 54 4/8. Pot 2 : 47 4/8 : 47 4/8. Pot 2 : 0 : 32 5/8.

Pot 3 : 54 7/8 : 46 5/8.

Pot 4 : 32 1/8 : 41 3/8. Pot 4 : 0 : 29 7/8. Pot 4 : 43 7/8 : 37 1/8.

Pot 5 : 46 6/8 : 42 1/8. Pot 5 : 40 4/8 : 42 1/8. Pot 5 : 43 : 0.

Pot 6 : 48 2/8 : 47 7/8. Pot 6 : 46 2/8 : 48 3/8.

Pot 7 : 48 5/8 : 25. Pot 7 : 42 : 40 5/8.

Pot 8 : 46 7/8 : 39 1/8.

Pot 9 : 49 : 30 3/8. Pot 9 : 50 3/8 : 15. Pot 9 : 46 3/8 : 36 7/8. Pot 9 : 47 6/8 : 44 1/8. Pot 9 : 0 : 31 6/8. Crowded Plants.

Pot 10 : 46 4/8 : 47 7/8. Pot 10 : 35 2/8 : 0. Pot 10 : 24 5/8 : 34 7/8. Pot 10 : 41 4/8 : 40 7/8. Pot 10 : 17 3/8 : 41 1/8. Crowded Plants.

Total : 1078.00 : 995.38.

The average height of the flower-stems on the twenty-five crossed plants in all the pots taken together is 43.12 inches, and that of the twenty-five self-fertilised plants 39.82, or as 100 to 92. In order to test this result, the plants planted in pairs in Pots 1 and 8 were considered by themselves, and the average height of the sixteen crossed plants is here 44.9, and that of the sixteen self-fertilised plants 42.03, or as 100 to 94. Again, the plants raised from the thickly sown seed in Pots 9 and 10, which were subjected to very severe mutual competition, were taken by themselves, and the average height of the nine crossed plants is 39.86, and that of the nine self-fertilised plants 35.88, or as 100 to 90. The plants in these two latter pots (9 and 10), after being measured, were cut down close to the ground and weighed: the nine crossed plants weighed 57.66 ounces, and the nine self-fertilised plants 45.25 ounces, or as 100 to 78. On the whole we may conclude, especially from the evidence of weight, that seedlings from a cross between flowers on the same plant have a decided, though not great, advantage over those from flowers fertilised with their own pollen, more especially in the case of the plants subjected to severe mutual competition. But the advantage is much less than that exhibited by the crossed offspring of distinct plants, for these exceeded the self-fertilised plants in height as 100 to 70, and in the number of flower-stems as 100 to 48. Digitalis thus differs from Ipomoea, and almost certainly from Mimulus, as with these two species a cross between flowers on the same plant did no good.

CALCEOLARIA.

A BUSHY GREENHOUSE VARIETY, WITH YELLOW FLOWERS BLOTCHED WITH PURPLE.

The flowers in this genus are constructed so as to favour or almost ensure cross-fertilisation (3/6. Hildebrand as quoted by H. Muller ‘Die Befruchtung der Blumen’ 1873 page 277.); and Mr. Anderson remarks that extreme care is necessary to exclude insects in order to preserve any kind true. (3/7. ‘Gardeners’ Chronicle’ 1853 page 534.) He adds the interesting statement, that when the corolla is cut quite away, insects, as far as he has seen, never discover or visit the flowers. This plant is, however, self-fertile if insects are excluded. So few experiments were made by me, that they are hardly worth giving. Crossed and self-fertilised seeds were sown on opposite sides of a pot, and after a time the crossed seedlings slightly exceeded the self-fertilised in height. When a little further grown, the longest leaves on the former were very nearly 3 inches in length, whilst those on the self-fertilised plants were only 2 inches. Owing to an accident, and to the pot being too small, only one plant on each side grew up and flowered; the crossed plant was 19 1/2 inches in height, and the self-fertilised one 15 inches; or as 100 to 77.

Linaria vulgaris.

It has been mentioned in the introductory chapter that two large beds of this plant were raised by me many years ago from crossed and self-fertilised seeds, and that there was a conspicuous difference in height and general appearance between the two lots. The trial was afterwards repeated with more care; but as this was one of the first plants experimented on, my usual method was not followed. Seeds were taken from wild plants growing in this neighbourhood and sown in poor soil in my garden. Five plants were covered with a net, the others being left exposed to the bees, which incessantly visit the flowers of this species, and which, according to H. Muller, are the exclusive fertilisers. This excellent observer remarks that, as the stigma lies between the anthers and is mature at the same time with them, self-fertilisation is possible. (3/8. ‘Die Befruchtung’ etc. page 279.) But so few seeds are produced by protected plants, that the pollen and stigma of the same flower seem to have little power of mutual interaction. The exposed plants bore numerous capsules forming solid spikes. Five of these capsules were examined and appeared to contain an equal number of seeds; and these being counted in one capsule, were found to be 166. The five protected plants produced altogether only twenty-five capsules, of which five were much finer than all the others, and these contained an average of 23.6 seeds, with a maximum in one capsule of fifty-five. So that the number of seeds in the capsules on the exposed plants to the average number in the finest capsules on the protected plants was as 100 to 14.

Some of the spontaneously self-fertilised seeds from under the net, and some seeds from the uncovered plants naturally fertilised and almost certainly intercrossed by the bees, were sown separately in two large pots of the same size; so that the two lots of seedlings were not subjected to any mutual competition. Three of the crossed plants when in full flower were measured, but no care was taken to select the tallest plants; their heights were 7 4/8, 7 2/8, and 6 4/8 inches; averaging 7.08 in height. The three tallest of all the self-fertilised plants were then carefully selected, and their heights were 6 3/8, 5 5/8, and 5 2/8, averaging 5.75 in height. So that the naturally crossed plants were to the spontaneously self-fertilised plants in height, at least as much as 100 to 81.

Verbascum thapsus.

The flowers of this plant are frequented by various insects, chiefly by bees, for the sake of the pollen. Hermann Muller, however, has shown (‘Die Befruchtung’ etc. page 277) that V. nigrum secretes minute drops of nectar. The arrangement of the reproductive organs, though not at all complex, favours cross-fertilisation; and even distinct species are often crossed, for a greater number of naturally produced hybrids have been observed in this genus than in almost any other. (3/9. I have given a striking case of a large number of such hybrids between Verbascum thapsus and lychnitis found growing wild: ‘Journal of Linnean Society Botany’ volume 10 page 451.) Nevertheless the present species is perfectly self-fertile, if insects are excluded; for a plant protected by a net was as thickly loaded with fine capsules as the surrounding uncovered plants. Verbascum lychnitis is rather less self-fertile, for some protected plants did not yield quite so many capsules as the adjoining uncovered plants.

Plants of Verbascum thapsus had been raised for a distinct purpose from self-fertilised seeds; and some flowers on these plants were again self-fertilised, yielding seed of the second self-fertilised generation; and other flowers were crossed with pollen from a distinct plant. The seeds thus produced were sown on the opposite sides of four large pots. They germinated, however, so irregularly (the crossed seedlings generally coming up first) that I was able to save only six pairs of equal age. These when in full flower were measured, as in Table 3/25.

TABLE 3/25. Verbascum thapsus.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants of the Second Generation.

Pot 1 : 76 : 53 4/8.

Pot 2 : 54 : 66.

Pot 3 : 62 : 75. Pot 3 : 60 5/8 : 30 4/8.

Pot 4 : 73 : 62. Pot 4 : 66 4/8 : 52.

Total : 392.13 : 339.00.

We here see that two of the self-fertilised plants exceed in height their crossed opponents. Nevertheless the average height of the six crossed plants is 65.34 inches, and that of the six self-fertilised plants 56.5 inches; or as 100 to 86.

Vandellia nummularifolia.

Seeds were sent to me by Mr. J. Scott from Calcutta of this small Indian weed, which bears perfect and cleistogene flowers. (3/10. The convenient term of CLEISTOGENE was proposed by Kuhn in an article on the present genus in ‘Bot. Zeitung’ 1867 page 65.) The latter are extremely small, imperfectly developed, and never expand, yet yield plenty of seeds. The perfect and open flowers are also small, of a white colour with purple marks; they generally produce seed, although the contrary has been asserted; and they do so even if protected from insects. They have a rather complicated structure, and appear to be adapted for cross-fertilisation, but were not carefully examined by me. They are not easy to fertilise artificially, and it is possible that some of the flowers which I thought that I had succeeded in crossing were afterwards spontaneously self-fertilised under the net. Sixteen capsules from the crossed perfect flowers contained on an average ninety-three seeds (with a maximum in one capsule of 137), and thirteen capsules from the self-fertilised perfect flowers contained sixty-two seeds (with a maximum in one capsule of 135); or as 100 to 67. But I suspect that this considerable excess was accidental, as on one occasion nine crossed capsules were compared with seven self-fertilised capsules (both included in the above number), and they contained almost exactly the same average number of seed. I may add that fifteen capsules from self-fertilised cleistogene flowers contained on an average sixty-four seeds, with a maximum in one of eighty-seven.

Crossed and self-fertilised seeds from the perfect flowers, and other seeds from the self-fertilised cleistogene flowers, were sown in five pots, each divided superficially into three compartments. The seedlings were thinned at an early age, so that twenty plants were left in each of the three divisions. The crossed plants when in full flower averaged 4.3 inches, and the self-fertilised plants from the perfect flowers 4.27 inches in height; or as 100 to 99. The self-fertilised plants from the cleistogene flowers averaged 4.06 inches in height; so that the crossed were in height to these latter plants as 100 to 94.

I determined to compare again the growth of plants raised from crossed and self-fertilised perfect flowers, and obtained two fresh lots of seeds. These were sown on opposite sides of five pots, but they were not sufficiently thinned, so that they grew rather crowded. When fully grown, all those above 2 inches in height were selected, all below this standard being rejected; the former consisted of forty-seven crossed and forty-one self-fertilised plants; thus a greater number of the crossed than of the self-fertilised plants grew to a height of above 2 inches. Of the crossed plants, the twenty-four tallest were on an average 3.6 inches in height; whilst the twenty-four tallest self-fertilised plants were 3.38 inches in average height; or as 100 to 94. All these plants were then cut down close to the ground, and the forty-seven crossed plants weighed 1090.3 grains, and the forty-one self-fertilised plants weighed 887.4 grains. Therefore an equal number of crossed and self-fertilised would have been to each other in weight as 100 to 97. From these several facts we may conclude that the crossed plants had some real, though very slight, advantage in height and weight over the self-fertilised plants, when grown in competition with one another.

The crossed plants were, however, inferior in fertility to the self-fertilised. Six of the finest plants were selected out of the forty-seven crossed plants, and six out of the forty-one self-fertilised plants; and the former produced 598 capsules, whilst the latter or self-fertilised plants produced 752 capsules. All these capsules were the product of cleistogene flowers, for the plants did not bear during the whole of this season any perfect flowers. The seeds were counted in ten cleistogene capsules produced by crossed plants, and their average number was 46.4 per capsule; whilst the number in ten cleistogene capsules produced by the self-fertilised plants was 49.4; or as 100 to 106.

3. GESNERIACEAE.—Gesneria pendulina.

In Gesneria the several parts of the flower are arranged on nearly the same plan as in Digitalis, and most or all of the species are dichogamous. (3/11. Dr. Ogle ‘Popular Science Review’ January 1870 page 51.) Plants were raised from seed sent me by Fritz Muller from South Brazil. Seven flowers were crossed with pollen from a distinct plant, and produced seven capsules containing by weight 3.01 grains of seeds. Seven flowers on the same plants were fertilised with their own pollen, and their seven capsules contained exactly the same weight of seeds. Germinating seeds were planted on opposite sides of four pots, and when fully grown measured to the tips of their leaves.

TABLE 3/26. Gesneria pendulina.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 42 2/8 : 39. Pot 1 : 24 4/8 : 27 3/8.

Pot 2 : 33 : 30 6/8. Pot 2 : 27 : 19 2/8.

Pot 3 : 33 4/8 : 31 7/8. Pot 3 : 29 4/8 : 28 6/8.

Pot 4 : 30 6/8 : 29 6/8. Pot 4 : 36 : 26 3/8.

Total : 256.50 : 233.13.

The average height of the eight crossed plants is 32.06 inches, and that of the eight self-fertilised plants 29.14; or as 100 to 90.

4. LABIATAE.—Salvia coccinea. (3/12. The admirable mechanical adaptations in this genus for favouring or ensuring cross-fertilisation, have been fully described by Sprengel, Hildebrand, Delpino, H. Muller, Ogle, and others, in their several works.)

This species, unlike most of the others in the same genus, yields a good many seeds when insects are excluded. I gathered ninety-eight capsules produced by flowers spontaneously self-fertilised under a net, and they contained on an average 1.45 seeds, whilst flowers artificially fertilised with their own pollen, in which case the stigma will have received plenty of pollen, yielded on an average 3.3 seeds, or more than twice as many. Twenty flowers were crossed with pollen from a distinct plant, and twenty-six were self-fertilised. There was no great difference in the proportional number of flowers which produced capsules by these two processes, or in the number of the contained seeds, or in the weight of an equal number of seeds.

Seeds of both kinds were sown rather thickly on opposite sides of three pots. When the seedlings were about 3 inches in height, the crossed showed a slight advantage over the self-fertilised. When two-thirds grown, the two tallest plants on each side of each pot were measured; the crossed averaged 16.37 inches, and the self-fertilised 11.75 in height; or as 100 to 71. When the plants were fully grown and had done flowering, the two tallest plants on each side were again measured, with the results shown in Table 3/27.

TABLE 3/27. Salvia coccinea.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 32 6/8 : 25. Pot 1 : 20 : 18 6/8.

Pot 2 : 32 3/8 : 20 6/8. Pot 2 : 24 4/8 : 19 4/8.

Pot 3 : 29 4/8 : 25. Pot 3 : 28 : 18.

Total : 167.13 : 127.00.

It may be here seen that each of the six tallest crossed plants exceeds in height its self-fertilised opponent; the former averaged 27.85 inches, whilst the six tallest self-fertilised plants averaged 21.16 inches; or as 100 to 76. In all three pots the first plant which flowered was a crossed one. All the crossed plants together produced 409 flowers, whilst all the self-fertilised together produced only 232 flowers; or as 100 to 57. So that the crossed plants in this respect were far more productive than the self-fertilised.

Origanum vulgare.

This plant exists, according to H. Muller, under two forms; one hermaphrodite and strongly proterandrous, so that it is almost certain to be fertilised by pollen from another flower; the other form is exclusively female, has a smaller corolla, and must of course be fertilised by pollen from a distinct plant in order to yield any seeds. The plants on which I experimented were hermaphrodites; they had been cultivated for a long period as a pot-herb in my kitchen garden, and were, like so many long-cultivated plants, extremely sterile. As I felt doubtful about the specific name I sent specimens to Kew, and was assured that the species was Origanum vulgare. My plants formed one great clump, and had evidently spread from a single root by stolons. In a strict sense, therefore, they all belonged to the same individual. My object in experimenting on them was, firstly, to ascertain whether crossing flowers borne by plants having distinct roots, but all derived asexually from the same individual, would be in any respect more advantageous than self-fertilisation; and, secondly, to raise for future trial seedlings which would constitute really distinct individuals. Several plants in the above clump were covered by a net, and about two dozen seeds (many of which, however, were small and withered) were obtained from the flowers thus spontaneously self-fertilised. The remainder of the plants were left uncovered and were incessantly visited by bees, so that they were doubtless crossed by them. These exposed plants yielded rather more and finer seed (but still very few) than did the covered plants. The two lots of seeds thus obtained were sown on opposite sides of two pots; the seedlings were carefully observed from their first growth to maturity, but they did not differ at any period in height or in vigour, the importance of which latter observation we shall presently see. When fully grown, the tallest crossed plant in one pot was a very little taller than the tallest self-fertilised plant on the opposite side, and in the other pot exactly the reverse occurred. So that the two lots were in fact equal; and a cross of this kind did no more good than crossing two flowers on the same plant of Ipomoea or Mimulus.

The plants were turned out of the two pots without being disturbed and planted in the open ground, in order that they might grow more vigorously. In the following summer all the self-fertilised and some of the quasi-crossed plants were covered by a net. Many flowers on the latter were crossed by me with pollen from a distinct plant, and others were left to be crossed by the bees. These quasi-crossed plants produced rather more seed than did the original ones in the great clump when left to the action of the bees. Many flowers on the self-fertilised plants were artificially self-fertilised, and others were allowed to fertilise themselves spontaneously under the net, but they yielded altogether very few seeds. These two lots of seeds—the product of a cross between distinct seedlings, instead of as in the last case between plants multiplied by stolons, and the product of self-fertilised flowers—were allowed to germinate on bare sand, and several equal pairs were planted on opposite sides of two LARGE pots. At a very early age the crossed plants showed some superiority over the self-fertilised, which was ever afterwards retained. When the plants were fully grown, the two tallest crossed and the two tallest self-fertilised plants in each pot were measured, as shown in Table 3/28. I regret that from want of time I did not measure all the pairs; but the tallest on each side seemed fairly to represent the average difference between the two lots.

TABLE 3/28. Origanum vulgare.

Heights of Plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants (two tallest in each pot).

Column 3: Self-fertilised Plants (two tallest in each pot).

Pot 1 : 26 : 24. Pot 1 : 21 : 21.

Pot 2 : 17 : 12. Pot 2 : 16 : 11 4/8.

Total : 80.0 : 68.5.

The average height of the crossed plants is here 20 inches, and that of the self-fertilised 17.12; or as 100 to 86. But this excess of height by no means gives a fair idea of the vast superiority in vigour of the crossed over the self-fertilised plants. The crossed flowered first and produced thirty flower-stems, whilst the self-fertilised produced only fifteen, or half the number. The pots were then bedded out, and the roots probably came out of the holes at the bottom and thus aided their growth. Early in the following summer the superiority of the crossed plants, owing to their increase by stolons, over the self-fertilised plants was truly wonderful. In Pot 1, and it should be remembered that very large pots had been used, the oval clump of crossed plants was 10 by 4 1/2 inches across, with the tallest stem, as yet young, 5 1/2 inches in height; whilst the clump of self-fertilised plants, on the opposite side of the same pot, was only 3 1/2 by 2 1/2 inches across, with the tallest young stem 4 inches in height. In Pot 2, the clump of crossed plants was 18 by 9 inches across, with the tallest young stem 8 1/2 inches in height; whilst the clump of self-fertilised plants on the opposite side of the same pot was 12 by 4 1/2 inches across, with the tallest young stem 6 inches in height. The crossed plants during this season, as during the last, flowered first. Both the crossed and self-fertilised plants being left freely exposed to the visits of bees, manifestly produced much more seed than their grand-parents,—the plants of the original clump still growing close by in the same garden, and equally left to the action of the bees.

5. ACANTHACEAE.—Thunbergia alata.

It appears from Hildebrand’s description (‘Botanische Zeitung’ 1867 page 285) that the conspicuous flowers of this plant are adapted for cross-fertilisation. Seedlings were twice raised from purchased seed; but during the early summer, when first experimented on, they were extremely sterile, many of the anthers containing hardly any pollen. Nevertheless, during the autumn these same plants spontaneously produced a good many seeds. Twenty-six flowers during the two years were crossed with pollen from a distinct plant, but they yielded only eleven capsules; and these contained very few seeds! Twenty-eight flowers were fertilised with pollen from the same flower, and these yielded only ten capsules, which, however, contained rather more seed than the crossed capsules. Eight pairs of germinating seeds were planted on opposite sides of five pots; and exactly half the crossed and half the self-fertilised plants exceeded their opponents in height. Two of the self-fertilised plants died young, before they were measured, and their crossed opponents were thrown away. The six remaining pairs of these grew very unequally, some, both of the crossed and self-fertilised plants, being more than twice as tall as the others. The average height of the crossed plants was 60 inches, and that of the self-fertilised plants 65 inches, or as 100 to 108. A cross, therefore, between distinct individuals here appears to do no good; but this result deduced from so few plants in a very sterile condition and growing very unequally, obviously cannot be trusted.]

CHAPTER IV. CRUCIFERAE, PAPAVERACEAE, RESEDACEAE, ETC.

Brassica oleracea, crossed and self-fertilised plants.
Great effect of a cross with a fresh stock on the weight of the
offspring.
Iberis umbellata.
Papaver vagum.
Eschscholtzia californica, seedlings from a cross with a fresh stock not
more vigorous, but more fertile than the self-fertilised seedlings.
Reseda lutea and odorata, many individuals sterile with their own pollen.
Viola tricolor, wonderful effects of a cross.
Adonis aestivalis.
Delphinium consolida.
Viscaria oculata, crossed plants hardly taller, but more fertile than
the self-fertilised.
Dianthus caryophyllus, crossed and self-fertilised plants compared for
four generations.
Great effects of a cross with a fresh stock.
Uniform colour of the flowers on the self-fertilised plants.
Hibiscus africanus.

[6. CRUCIFERAE.—Brassica oleracea.

VAR. CATTELL’S EARLY BARNES CABBAGE.

The flowers of the common cabbage are adapted, as shown by H. Muller, for cross-fertilisation, and should this fail, for self-fertilisation. (4/1. ‘Die Befruchtung’ etc. page 139.) It is well known that the varieties are crossed so largely by insects, that it is impossible to raise pure kinds in the same garden, if more than one kind is in flower at the same time. Cabbages, in one respect, were not well fitted for my experiments, as, after they had formed heads, they were often difficult to measure. The flower-stems also differ much in height; and a poor plant will sometimes throw up a higher stem than that of a fine plant. In the later experiments, the fully-grown plants were cut down and weighed, and then the immense advantage from a cross became manifest.

A single plant of the above variety was covered with a net just before flowering, and was crossed with pollen from another plant of the same variety growing close by; and the seven capsules thus produced contained on an average 16.3 seeds, with a maximum of twenty in one capsule. Some flowers were artificially self-fertilised, but their capsules did not contain so many seeds as those from flowers spontaneously self-fertilised under the net, of which a considerable number were produced. Fourteen of these latter capsules contained on an average 4.1 seeds, with a maximum in one of ten seeds; so that the seeds in the crossed capsules were in number to those in the self-fertilised capsules as 100 to 25. The self-fertilised seeds, fifty-eight of which weighed 3.88 grains, were, however, a little finer than those from the crossed capsules, fifty-eight of which weighed 3.76 grains. When few seeds are produced, these seem often to be better nourished and to be heavier than when many are produced.

The two lots of seeds in an equal state of germination were planted, some on opposite sides of a single pot, and some in the open ground. The young crossed plants in the pot at first exceeded by a little in height the self-fertilised; then equalled them; were then beaten; and lastly were again victorious. The plants, without being disturbed, were turned out of the pot, and planted in the open ground; and after growing for some time, the crossed plants, which were all of nearly the same height, exceeded the self-fertilised ones by 2 inches. When they flowered, the flower-stems of the tallest crossed plant exceeded that of the tallest self-fertilised plant by 6 inches. The other seedlings which were planted in the open ground stood separate, so that they did not compete with one another; nevertheless the crossed plants certainly grew to a rather greater height than the self-fertilised; but no measurements were made. The crossed plants which had been raised in the pot, and those planted in the open ground, all flowered a little before the self-fertilised plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

Some flowers on the crossed plants of the last generation were again crossed with pollen from another crossed plant, and produced fine capsules. The flowers on the self-fertilised plants of the last generation were allowed to fertilise themselves spontaneously under a net, and they produced some remarkably fine capsules. The two lots of seeds thus produced germinated on sand, and eight pairs were planted on opposite sides of four pots. These plants were measured to the tips of their leaves on the 20th of October of the same year, and the eight crossed plants averaged in height 8.4 inches, whilst the self-fertilised averaged 8.53 inches, so that the crossed were a little inferior in height, as 100 to 101.5. By the 5th of June of the following year these plants had grown much bulkier, and had begun to form heads. The crossed had now acquired a marked superiority in general appearance, and averaged 8.02 inches in height, whilst the self-fertilised averaged 7.31 inches; or as 100 to 91. The plants were then turned out of their pots and planted undisturbed in the open ground. By the 5th of August their heads were fully formed, but several had grown so crooked that their heights could hardly be measured with accuracy. The crossed plants, however, were on the whole considerably taller than the self-fertilised. In the following year they flowered; the crossed plants flowering before the self-fertilised in three of the pots, and at the same time in Pot 2. The flower-stems were now measured, as shown in Table 4/29.

TABLE 3/29. Brassica oleracea.

Measured in inches to tops of flower-stems: 0 signifies that a Flower-stem was not formed.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 49 2/8 : 44. Pot 1 : 39 4/8 : 41.

Pot 2 : 37 4/8 : 38. Pot 2 : 33 4/8 : 35 4/8.

Pot 3 : 47 : 51 1/8. Pot 3 : 40 : 41 2/8. Pot 3 : 42 : 46 4/8.

Pot 4 : 43 6/8 : 20 2/8. Pot 4 : 37 2/8 : 33 3/8. Pot 4 : 0 : 0.

Total : 369.75 : 351.00.

The nine flower-stems on the crossed plants here average 41.08 inches, and the nine on the self-fertilised plants 39 inches in height, or as 100 to 95. But this small difference, which, moreover, depended almost wholly on one of the self-fertilised plants being only 20 inches high, does not in the least show the vast superiority of the crossed over the self-fertilised plants. Both lots, including the two plants in Pot 4, which did not flower, were now cut down close to the ground and weighed, but those in Pot 2 were excluded, for they had been accidentally injured by a fall during transplantation, and one was almost killed. The eight crossed plants weighed 219 ounces, whilst the eight self-fertilised plants weighed only 82 ounces, or as 100 to 37; so that the superiority of the former over the latter in weight was great.

THE EFFECTS OF A CROSS WITH A FRESH STOCK.

Some flowers on a crossed plant of the last or second generation were fertilised, without being castrated, by pollen taken from a plant of the same variety, but not related to my plants, and brought from a nursery garden (whence my seeds originally came) having a different soil and aspect. The flowers on the self-fertilised plants of the last or second generation (Table 4/29) were allowed to fertilise themselves spontaneously under a net, and yielded plenty of seeds. These latter and the crossed seeds, after germinating on sand, were planted in pairs on the opposite sides of six large pots, which were kept at first in a cool greenhouse. Early in January their heights were measured to the tips of their leaves. The thirteen crossed plants averaged 13.16 inches in height, and the twelve (for one had died) self-fertilised plants averaged 13.7 inches, or as 100 to 104; so that the self-fertilised plants exceeded by a little the crossed plants.

TABLE 3/30. Brassica oleracea.

Weights in ounces of plants after they had formed heads.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants from Pollen of fresh Stock.

Column 3: Self-fertilised Plants of the Third Generation.

Pot 1 : 130 : 18 2/4.

Pot 2 : 74 : 34 3/4.

Pot 3 : 121 : 17 2/4.

Pot 4 : 127 2/4 : 14.

Pot 5 : 90 : 11 2/4.

Pot 6 : 106 2/4 : 46.

Total : 649.00 : 142.25.

Early in the spring the plants were gradually hardened, and turned out of their pots into the open ground without being disturbed. By the end of August the greater number had formed fine heads, but several grew extremely crooked, from having been drawn up to the light whilst in the greenhouse. As it was scarcely possible to measure their heights, the finest plant on each side of each pot was cut down close to the ground and weighed. In Table 4/30 we have the result.

The six finest crossed plants average 108.16 ounces, whilst the six finest self-fertilised plants average only 23.7 ounces, or as 100 to 22. This difference shows in the clearest manner the enormous benefit which these plants derived from a cross with another plant belonging to the same sub-variety, but to a fresh stock, and grown during at least the three previous generations under somewhat different conditions.

THE OFFSPRING FROM A CUT-LEAVED, CURLED, AND VARIEGATED WHITE-GREEN CABBAGE CROSSED WITH A CUT-LEAVED, CURLED, AND VARIEGATED CRIMSON-GREEN CABBAGE, COMPARED WITH THE SELF-FERTILISED OFFSPRING FROM THE TWO VARIETIES.

These trials were made, not for the sake of comparing the growth of the crossed and self-fertilised seedlings, but because I had seen it stated that these varieties would not naturally intercross when growing uncovered and near one another. This statement proved quite erroneous; but the white-green variety was in some degree sterile in my garden, producing little pollen and few seeds. It was therefore no wonder that seedlings raised from the self-fertilised flowers of this variety were greatly exceeded in height by seedlings from a cross between it and the more vigorous crimson-green variety; and nothing more need be said about this experiment.

The seedlings from the reciprocal cross, that is, from the crimson-green variety fertilised with pollen from the white-green variety, offer a somewhat more curious case. A few of these crossed seedlings reverted to a pure green variety with their leaves less cut and curled, so that they were altogether in a much more natural state, and these plants grew more vigorously and taller than any of the others. Now it is a strange fact that a much larger number of the self-fertilised seedlings from the crimson-green variety than of the crossed seedlings thus reverted; and as a consequence the self-fertilised seedlings grew taller by 2 1/2 inches on an average than the crossed seedlings, with which they were put into competition. At first, however, the crossed seedlings exceeded the self-fertilised by an average of a quarter of an inch. We thus see that reversion to a more natural condition acted more powerfully in favouring the ultimate growth of these plants than did a cross; but it should be remembered that the cross was with a semi-sterile variety having a feeble constitution.

Iberis umbellata.

VAR. KERMESIANA.

This variety produced plenty of spontaneously self-fertilised seed under a net. Other plants in pots in the greenhouse were left uncovered, and as I saw small flies visiting the flowers, it seemed probable that they would be intercrossed. Consequently seeds supposed to have been thus crossed and spontaneously self-fertilised seeds were sown on opposite sides of a pot. The self-fertilised seedlings grew from the first quicker than the supposed crossed seedlings, and when both lots were in full flower the former were from 5 to 6 inches higher than the crossed! I record in my notes that the self-fertilised seeds from which these self-fertilised plants were raised were not so well ripened as the crossed; and this may possibly have caused the great difference in their growth, in a somewhat analogous manner as occurred with the self-fertilised plants of the eighth generation of Ipomoea raised from unhealthy parents. It is a curious circumstance, that two other lots of the above seeds were sown in pure sand mixed with burnt earth, and therefore without any organic matter; and here the supposed crossed seedlings grew to double the height of the self-fertilised, before both lots died, as necessarily occurred at an early period. We shall hereafter meet with another case apparently analogous to this of Iberis in the third generation of Petunia.

The above self-fertilised plants were allowed to fertilise themselves again under a net, yielding self-fertilised plants of the second generation, and the supposed crossed plants were crossed by pollen of a distinct plant; but from want of time this was done in a careless manner, namely, by smearing one head of expanded flowers over another. I should have thought that this would have succeeded, and perhaps it did so; but the fact of 108 of the self-fertilised seeds weighing 4.87 grains, whilst the same number of the supposed crossed seeds weighed only 3.57 grains, does not look like it. Five seedlings from each lot of seeds were raised, and the self-fertilised plants, when fully grown, exceeded in average height by a trifle (namely .4 of an inch) the five probably crossed plants. I have thought it right to give this case and the last, because had the supposed crossed plants proved superior to the self-fertilised in height, I should have assumed without doubt that the former had really been crossed. As it is, I do not know what to conclude.

Being much surprised at the two foregoing trials, I determined to make another, in which there should be no doubt about the crossing. I therefore fertilised with great care (but as usual without castration) twenty-four flowers on the supposed crossed plants of the last generation with pollen from distinct plants, and thus obtained twenty-one capsules. The self-fertilised plants of the last generation were allowed to fertilise themselves again under a net, and the seedlings reared from these seeds formed the third self-fertilised generation. Both lots of seeds, after germinating on bare sand, were planted in pairs on the opposite sides of two pots. All the remaining seeds were sown crowded on opposite sides of a third pot; but as all the self-fertilised seedlings in this latter pot died before they grew to any considerable height, they were not measured. The plants in Pots 1 and 2 were measured when between 7 and 8 inches in height, and the crossed exceeded the self-fertilised in average height by 1.57 inches. When fully grown they were again measured to the summits of their flower-heads, with the following result:—

TABLE 4/31. Iberis umbellata.

Heights of plants to the summits of their flower-heads, in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants of the Third Generation.

Pot 1 : 18 : 19. Pot 1 : 21 : 21. Pot 1 : 18 2/8 : 19 4/8.

Pot 2 : 19 : 16 6/8. Pot 2 : 18 4/8 : 7 4/8. Pot 2 : 17 6/8 : 14 4/8. Pot 2 : 21 3/8 : 16 4/8.

Total : 133.88 : 114.75.

The average height of the seven crossed plants is here 19.12 inches, and that of the seven self-fertilised plants 16.39, or as 100 to 86. But as the plants on the self-fertilised side grew very unequally, this ratio cannot be fully trusted, and is probably too high. In both pots a crossed plant flowered before any one of the self-fertilised. These plants were left uncovered in the greenhouse; but from being too much crowded they were not very productive. The seeds from all seven plants of both lots were counted; the crossed produced 206, and the self-fertilised 154; or as 100 to 75.

CROSS BY A FRESH STOCK.

From the doubts caused by the two first trials, in which it was not known with certainty that the plants had been crossed; and from the crossed plants in the last experiment having been put into competition with plants self-fertilised for three generations, which moreover grew very unequally, I resolved to repeat the trial on a larger scale, and in a rather different manner. I obtained seeds of the same crimson variety of Iberis umbellata from another nursery garden, and raised plants from them. Some of these plants were allowed to fertilise themselves spontaneously under a net; others were crossed by pollen taken from plants raised from seed sent me by Dr. Durando from Algiers, where the parent-plants had been cultivated for some generations. These latter plants differed in having pale pink instead of crimson flowers, but in no other respect. That the cross had been effective (though the flowers on the crimson mother-plant had NOT been castrated) was well shown when the thirty crossed seedlings flowered, for twenty-four of them produced pale pink flowers, exactly like those of their father; the six others having crimson flowers exactly like those of their mother and like those of all the self-fertilised seedlings. This case offers a good instance of a result which not rarely follows from crossing varieties having differently coloured flowers; namely, that the colours do not blend, but resemble perfectly those either of the father or mother plant. The seeds of both lots, after germinating on sand, were planted on opposite sides of eight pots. When fully grown, the plants were measured to the summits of the flower-heads, as shown in Table 4/32.

TABLE 4/32. Iberis umbellata.

Height of Plants to the summits of the flower-heads, measured in inches: 0 signifies that the Plant died.

Column 1: Number (Name) of Pot.

Column 2: Plants from a Cross with a fresh Stock.

Column 3: Plants from Spontaneously Self-fertilised Seeds.

Pot 1 : 18 6/8 : 17 3/8. Pot 1 : 17 5/8 : 16 7/8. Pot 1 : 17 6/8 : 13 1/8. Pot 1 : 20 1/8 : 15 3/8.

Pot 2 : 20 2/8 : 0. Pot 2 : 15 7/8 : 16 6/8. Pot 2 : 17 : 15 2/8.

Pot 3 : 19 2/8 : 13 6/8. Pot 3 : 18 1/8 : 14 2/8. Pot 3 : 15 2/8 : 13 4/8.

Pot 4 : 17 1/8 : 16 4/8. Pot 4 : 18 7/8 : 14 4/8. Pot 4 : 17 5/8 : 16. Pot 4 : 15 6/8 : 15 3/8. Pot 4 : 14 4/8 : 14 7/8.

Pot 5 : 18 1/8 : 16 4/8. Pot 5 : 14 7/8 : 16 2/8. Pot 5 : 16 2/8 : 14 2/8. Pot 5 : 15 5/8 : 14 2/8. Pot 5 : 12 4/8 : 16 1/8.

Pot 6 : 18 6/8 : 16 1/8. Pot 6 : 18 6/8 : 15. Pot 6 : 17 3/8 : 15 2/8.

Pot 7 : 18 : 16 3/8. Pot 7 : 16 4/8 : 14 4/8. Pot 7 : 18 2/8 : 13 5/8.

Pot 8 : 20 6/8 : 15 6/8. Pot 8 : 17 7/8 : 16 3/8. Pot 8 : 13 5/8 : 20 2/8. Pot 8 : 19 2/8 : 15 6/8.

Total : 520.38 : 449.88.

The average height of the thirty crossed plants is here 17.34, and that of the twenty-nine self-fertilised plants (one having died) 15.51, or as 100 to 89. I am surprised that the difference did not prove somewhat greater, considering that in the last experiment it was as 100 to 86; but this latter ratio, as before explained, was probably too great. It should, however, be observed that in the last experiment (Table 4/31), the crossed plants competed with plants of the third self-fertilised generation; whilst in the present case, plants derived from a cross with a fresh stock competed with self-fertilised plants of the first generation.

The crossed plants in the present case, as in the last, were more fertile than the self-fertilised, both lots being left uncovered in the greenhouse. The thirty crossed plants produced 103 seed-bearing flowers-heads, as well as some heads which yielded no seeds; whereas the twenty-nine self-fertilised plants produced only 81 seed-bearing heads; therefore thirty such plants would have produced 83.7 heads. We thus get the ratio of 100 to 81, for the number of seed-bearing flower-heads produced by the crossed and self-fertilised plants. Moreover, a number of seed-bearing heads from the crossed plants, compared with the same number from the self-fertilised, yielded seeds by weight, in the ratio of 100 to 92. Combining these two elements, namely, the number of seed-bearing heads and the weight of seeds in each head, the productiveness of the crossed to the self-fertilised plants was as 100 to 75.

The crossed and self-fertilised seeds, which remained after the above pairs had been planted, (some in a state of germination and some not so), were sown early in the year out of doors in two rows. Many of the self-fertilised seedlings suffered greatly, and a much larger number of them perished than of the crossed. In the autumn the surviving self-fertilised plants were plainly less well-grown than the crossed plants.

7. PAPAVERACEAE.—Papaver vagum.

A SUB-SPECIES OF Papaver dubium, FROM THE SOUTH OF FRANCE.

The poppy does not secrete nectar, but the flowers are highly conspicuous and are visited by many pollen-collecting bees, flies and beetles. The anthers shed their pollen very early, and in the case of Papaver rhoeas, it falls on the circumference of the radiating stigmas, so that this species must often be self-fertilised; but with Papaver dubium the same result does not follow (according to H. Muller ‘Die Befruchtung’ page 128), owing to the shortness of the stamens, unless the flower happens to stand inclined. The present species, therefore, does not seem so well fitted for self-fertilisation as most of the others. Nevertheless Papaver vagum produced plenty of capsules in my garden when insects were excluded, but only late in the season. I may here add that Papaver somniferum produces an abundance of spontaneously self-fertilised capsules, as Professor H. Hoffmann likewise found to be the case. (4/2. ‘Zur Speciesfrage’ 1875 page 53.) Some species of Papaver cross freely when growing in the same garden, as I have known to be the case with Papaver bracteatum and orientale.

Plants of Papaver vagum were raised from seeds sent me from Antibes through the kindness of Dr. Bornet. Some little time after the flowers had expanded, several were fertilised with their own pollen, and others (not castrated) with pollen from a distinct individual; but I have reason to believe, from observations subsequently made, that these flowers had been already fertilised by their own pollen, as this process seems to take place soon after their expansion. (4/3. Mr. J. Scott found ‘Report on the Experimental Culture of the Opium Poppy’ Calcutta 1874 page 47, in the case of Papaver somniferum, that if he cut away the stigmatic surface before the flower had expanded, no seeds were produced; but if this was done “on the second day, or even a few hours after the expansion of the flower on the first day, a partial fertilisation had already been effected, and a few good seeds were almost invariably produced.” This proves at how early a period fertilisation takes place.) I raised, however, a few seedlings of both lots, and the self-fertilised rather exceeded the crossed plants in height.

Early in the following year I acted differently, and fertilised seven flowers, very soon after their expansion, with pollen from another plant, and obtained six capsules. From counting the seeds in a medium-sized one, I estimated that the average number in each was at least 120. Four out of twelve capsules, spontaneously self-fertilised at the same time, were found to contain no good seeds; and the remaining eight contained on an average 6.6 seeds per capsule. But it should be observed that later in the season the same plants produced under a net plenty of very fine spontaneously self-fertilised capsules.

The above two lots of seeds, after germinating on sand, were planted in pairs on opposite sides of five pots. The two lots of seedlings, when half an inch in height, and again when 6 inches high, were measured to the tips of their leaves, but presented no difference. When fully grown, the flower-stalks were measured to the summits of the seed capsules, with the following result:—

TABLE 4/33. Papaver vagum.

Heights of flower-stalks to the summits of the seed capsules measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 24 2/8 : 21. Pot 1 : 30 : 26 5/8. Pot 1 : 18 4/8 : 16.

Pot 2 : 14 4/8 : 15 3/8. Pot 2 : 22 : 20 1/8. Pot 2 : 19 5/8 : 14 1/8. Pot 2 : 21 5/8 : 16 4/8.

Pot 3 : 20 6/8 : 19 2/8. Pot 3 : 20 2/8 : 13 2/8. Pot 3 : 20 6/8 : 18.

Pot 4 : 25 3/8 : 23 2/8. Pot 4 : 24 2/8 : 23.

Pot 5 : 20 : 18 3/8. Pot 5 : 27 7/8 : 27. Pot 5 : 19 : 21 2/8.

Total : 328.75 : 293.13.

The fifteen crossed plants here average 21.91 inches, and the fifteen self-fertilised plants 19.54 inches in height, or as 100 to 89. These plants did not differ in fertility, as far as could be judged by the number of capsules produced, for there were seventy-five on the crossed side and seventy-four on the self-fertilised side.

Eschscholtzia californica.

This plant is remarkable from the crossed seedlings not exceeding in height or vigour the self-fertilised. On the other hand, a cross greatly increases the productiveness of the flowers on the parent-plant, and is indeed sometimes necessary in order that they should produce any seed; moreover, plants thus derived are themselves much more fertile than those raised from self-fertilised flowers; so that the whole advantage of a cross is confined to the reproductive system. It will be necessary for me to give this singular case in considerable detail.

Twelve flowers on some plants in my flower-garden were fertilised with pollen from distinct plants, and produced twelve capsules; but one of these contained no good seed. The seeds of the eleven good capsules weighed 17.4 grains. Eighteen flowers on the same plants were fertilised with their own pollen and produced twelve good capsules, which contained 13.61 grains weight of seed. Therefore an equal number of crossed and self-fertilised capsules would have yielded seed by weight as 100 to 71. (4/4. Professor Hildebrand experimented on plants in Germany on a larger scale than I did, and found them much more self-fertile. Eighteen capsules, produced by cross-fertilisation, contained on an average eighty-five seeds, whilst fourteen capsules from self-fertilised flowers contained on an average only nine seeds; that is, as 100 to 11: ‘Jahrb. fur Wissen Botanik.’ B. 7 page 467.) If we take into account of the fact that a much greater proportion of flowers produced capsules when crossed than when self-fertilised, the relative fertility of the crossed to the self-fertilised flowers was as 100 to 52. Nevertheless these plants, whilst still protected by the net, spontaneously produced a considerable number of self-fertilised capsules.

The seeds of the two lots after germinating on sand were planted in pairs on the opposite sides of four large pots. At first there was no difference in their growth, but ultimately the crossed seedlings exceeded the self-fertilised considerably in height, as shown in Table 4/34. But I believe from the cases which follow that this result was accidental, owing to only a few plants having been measured, and to one of the self-fertilised plants having grown only to a height of 15 inches. The plants had been kept in the greenhouse, and from being drawn up to the light had to be tied to sticks in this and the following trials. They were measured to the summits of their flower-stems.

TABLE 4/34. Eschscholtzia californica.

Heights of Plants to the summits of their flower-stems measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 33 4/8 : 25.

Pot 2 : 34 2/8 : 35.

Pot 3 : 29 : 27 2/8.

Pot 4 : 22 : 15.

Total : 118.75 : 102.25.

The four crossed plants here average 29.68 inches, and the four self-fertilised 25.56 in height; or as 100 to 86. The remaining seeds were sown in a large pot in which a Cineraria had long been growing; and in this case again the two crossed plants on the one side greatly exceeded in height the two self-fertilised plants on the opposite side. The plants in the above four pots from having been kept in the greenhouse did not produce on this or any other similar occasion many capsules; but the flowers on the crossed plants when again crossed were much more productive than the flowers on the self-fertilised plants when again self-fertilised. These plants after seeding were cut down and kept in the greenhouse; and in the following year, when grown again, their relative heights were reversed, as the self-fertilised plants in three out of the four pots were now taller than and flowered before the crossed plants.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

The fact just given with respect to the growth of the cut-down plants made me doubtful about my first trial, so I determined to make another on a larger scale with crossed and self-fertilised seedlings raised from the crossed and self-fertilised plants of the last generation. Eleven pairs were raised and grown in competition in the usual manner; and now the result was different, for the two lots were nearly equal during their whole growth. It would therefore be superfluous to give a table of their heights. When fully grown and measured, the crossed averaged 32.47, and the self-fertilised 32.81 inches in height; or as 100 to 101. There was no great difference in the number of flowers and capsules produced by the two lots when both were left freely exposed to the visits of insects.

PLANTS RAISED FROM BRAZILIAN SEED.

Fritz Muller sent me from South Brazil seeds of plants which were there absolutely sterile when fertilised with pollen from the same plant, but were perfectly fertile when fertilised with pollen from any other plant. The plants raised by me in England from these seeds were examined by Professor Asa Gray, and pronounced to belong to E. Californica, with which they were identical in general appearance. Two of these plants were covered by a net, and were found not to be so completely self-sterile as in Brazil. But I shall recur to this subject in another part of this work. Here it will suffice to state that eight flowers on these two plants, fertilised with pollen from another plant under the net, produced eight fine capsules, each containing on an average about eighty seeds. Eight flowers on these same plants, fertilised with their own pollen, produced seven capsules, which contained on an average only twelve seeds, with a maximum in one of sixteen seeds. Therefore the cross-fertilised capsules, compared with the self-fertilised, yielded seeds in the ratio of about 100 to 15. These plants of Brazilian parentage differed also in a marked manner from the English plants in producing extremely few spontaneously self-fertilised capsules under a net.

Crossed and self-fertilised seeds from the above plants, after germinating on bare sand, were planted in pairs on the opposite sides of five large pots. The seedlings thus raised were the grandchildren of the plants which grew in Brazil; the parents having been grown in England. As the grandparents in Brazil absolutely require cross-fertilisation in order to yield any seeds, I expected that self-fertilisation would have proved very injurious to these seedlings, and that the crossed ones would have been greatly superior in height and vigour to those raised from self-fertilised flowers. But the result showed that my anticipation was erroneous; for as in the last experiment with plants of the English stock, so in the present one, the self-fertilised plants exceeded the crossed by a little in height. It will be sufficient to state that the fourteen crossed plants averaged 44.64, and the fourteen self-fertilised 45.12 inches in height; or as 100 to 101.

THE EFFECTS OF A CROSS WITH A FRESH STOCK.

I now tried a different experiment. Eight flowers on the self-fertilised plants of the last experiment (i.e., grandchildren of the plants which grew in Brazil) were again fertilised with pollen from the same plant, and produced five capsules, containing on an average 27.4 seeds, with a maximum in one of forty-two seeds. The seedlings raised from these seeds formed the second SELF-FERTILISED generation of the Brazilian stock.

Eight flowers on one of the crossed plants of the last experiment were crossed with pollen from another grandchild, and produced five capsules. These contained on an average 31.6 seeds, with a maximum in one of forty-nine seeds. The seedlings raised from these seeds may be called the INTERCROSSED.

Lastly, eight other flowers on the crossed plants of the last experiment were fertilised with pollen from a plant of the English stock, growing in my garden, and which must have been exposed during many previous generations to very different conditions from those to which the Brazilian progenitors of the mother-plant had been subjected. These eight flowers produced only four capsules, containing on an average 63.2 seeds, with a maximum in one of ninety. The plants raised from these seeds may be called the ENGLISH-CROSSED. As far as the above averages can be trusted from so few capsules, the English-crossed capsules contained twice as many seeds as the intercrossed, and rather more than twice as many as the self-fertilised capsules. The plants which yielded these capsules were grown in pots in the greenhouse, so that their absolute productiveness must not be compared with that of plants growing out of doors.

The above three lots of seeds, namely, the self-fertilised, intercrossed, and English-crossed, were planted in an equal state of germination (having been as usual sown on bare sand) in nine large pots, each divided into three parts by superficial partitions. Many of the self-fertilised seeds germinated before those of the two crossed lots, and these were of course rejected. The seedlings thus raised are the great-grandchildren of the plants which grew in Brazil. When they were from 2 to 4 inches in height, the three lots were equal. They were measured when four-fifths grown, and again when fully grown, and as their relative heights were almost exactly the same at these two ages, I will give only the last measurements. The average height of the nineteen English-crossed plants was 45.92 inches; that of the eighteen intercrossed plants (for one died), 43.38; and that of the nineteen self-fertilised plants, 50.3 inches. So that we have the following ratios in height:—

The English-crossed to the self-fertilised plants, as 100 to 109.

The English-crossed to the intercrossed plants, as 100 to 94.

The intercrossed to the self-fertilised plants, as 100 to 116.

After the seed-capsules had been gathered, all these plants were cut down close to the ground and weighed. The nineteen English crossed plants weighed 18.25 ounces; the intercrossed plants (with their weight calculated as if there had been nineteen) weighed 18.2 ounces; and the nineteen self-fertilised plants, 21.5 ounces. We have therefore for the weights of the three lots of plants the following ratios:—

The English-crossed to the self-fertilised plants, as 100 to 118.

The English-crossed to the intercrossed plants, as 100 to 100.

The intercrossed to the self-fertilised plants, as 100 to 118.

We thus see that in weight, as in height, the self-fertilised plants had a decided advantage over the English-crossed and intercrossed plants.

The remaining seeds of the three kinds, whether or not in a state of germination, were sown in three long parallel rows in the open ground; and here again the self-fertilised seedlings exceeded in height by between 2 and 3 inches the seedlings in the two other rows, which were of nearly equal heights. The three rows were left unprotected throughout the winter, and all the plants were killed, with the exception of two of the self-fertilised; so that as far as this little bit of evidence goes, some of the self-fertilised plants were more hardy than any of the crossed plants of either lot.

We thus see that the self-fertilised plants which were grown in the nine pots were superior in height (as 116 to 100), and in weight (as 118 to 100), and apparently in hardiness, to the intercrossed plants derived from a cross between the grandchildren of the Brazilian stock. The superiority is here much more strongly marked than in the second trial with the plants of the English stock, in which the self-fertilised were to the crossed in height as 101 to 100. It is a far more remarkable fact—if we bear in mind the effects of crossing plants with pollen from a fresh stock in the cases of Ipomoea, Mimulus, Brassica, and Iberis—that the self-fertilised plants exceeded in height (as 109 to 100), and in weight (as 118 to 100), the offspring of the Brazilian stock crossed by the English stock; the two stocks having been long subjected to widely different conditions.

If we now turn to the fertility of the three lots of plants we find a very different result. I may premise that in five out of the nine pots the first plant which flowered was one of the English-crossed; in four of the pots it was a self-fertilised plant; and in not one did an intercrossed plant flower first; so that these latter plants were beaten in this respect, as in so many other ways. The three closely adjoining rows of plants growing in the open ground flowered profusely, and the flowers were incessantly visited by bees, and certainly thus intercrossed. The manner in which several plants in the previous experiments continued to be almost sterile as long as they were covered by a net, but set a multitude of capsules immediately that they were uncovered, proves how effectually the bees carry pollen from plant to plant. My gardener gathered, at three successive times, an equal number of ripe capsules from the plants of the three lots, until he had collected forty-five from each lot. It is not possible to judge from external appearance whether or not a capsule contains any good seeds; so that I opened all the capsules. Of the forty-five from the English-crossed plants, four were empty; of those from the intercrossed, five were empty; and of those from the self-fertilised, nine were empty. The seeds were counted in twenty-one capsules taken by chance out of each lot, and the average number of seeds in the capsules from the English-crossed plants was 67; from the intercrossed, 56; and from the self-fertilised, 48.52. It therefore follows that:—

The forty-five capsules (the four empty ones included) from the English-crossed plants contained 2747 seeds.

The forty-five capsules (the five empty ones included) from the intercrossed plants contained 2240 seeds.

The forty-five capsules (the nine empty ones included) from the self-fertilised plants contained 1746.7 seeds.

The reader should remember that these capsules are the product of cross-fertilisation, effected by the bees; and that the difference in the number of the contained seeds must depend on the constitution of the plants;—that is, on whether they were derived from a cross with a distinct stock, or from a cross between plants of the same stock, or from self-fertilisation. From the above facts we obtain the following ratios:—

Number of seeds contained in an equal number of naturally fertilised capsules produced:—

By the English-crossed and self-fertilised plants, as 100 to 63.

By the English-crossed and intercrossed plants, as 100 to 81.

By the intercrossed and self-fertilised plants, as 100 to 78.

But to have ascertained the productiveness of the three lots of plants, it would have been necessary to know how many capsules were produced by the same number of plants. The three long rows, however, were not of quite equal lengths, and the plants were much crowded, so that it would have been extremely difficult to have ascertained how many capsules were produced by them, even if I had been willing to undertake so laborious a task as to collect and count all the capsules. But this was feasible with the plants grown in pots in the greenhouse; and although these were much less fertile than those growing out of doors, their relative fertility appeared, after carefully observing them, to be the same. The nineteen plants of the English-crossed stock in the pots produced altogether 240 capsules; the intercrossed plants (calculated as nineteen) produced 137.22 capsules; and the nineteen self-fertilised plants, 152 capsules. Now, knowing the number of seeds contained in forty-five capsules of each lot, it is easy to calculate the relative numbers of seeds produced by an equal number of the plants of the three lots.

Number of seeds produced by an equal number of naturally-fertilised plants:—

Plants of English-crossed and self-fertilised parentage, as 100 to 40 seeds.

Plants of English-crossed and intercrossed parentage, as 100 to 45 seeds.

Plants of intercrossed and self-fertilised parentage, as 100 to 89 seeds.

The superiority in productiveness of the intercrossed plants (that is, the product of a cross between the grandchildren of the plants which grew in Brazil) over the self-fertilised, small as it is, is wholly due to the larger average number of seeds contained in the capsules; for the intercrossed plants produced fewer capsules in the greenhouse than did the self-fertilised plants. The great superiority in productiveness of the English-crossed over the self-fertilised plants is shown by the larger number of capsules produced, the larger average number of contained seeds, and the smaller number of empty capsules. As the English-crossed and intercrossed plants were the offspring of crosses in every previous generation (as must have been the case from the flowers being sterile with their own pollen), we may conclude that the great superiority in productiveness of the English-crossed over the intercrossed plants is due to the two parents of the former having been long subjected to different conditions.

The English-crossed plants, though so superior in productiveness, were, as we have seen, decidedly inferior in height and weight to the self-fertilised, and only equal to, or hardly superior to, the intercrossed plants. Therefore, the whole advantage of a cross with a distinct stock is here confined to productiveness, and I have met with no similar case.

8. RESEDACEAE.—Reseda lutea.

Seeds collected from wild plants growing in this neighbourhood were sown in the kitchen-garden; and several of the seedlings thus raised were covered with a net. Of these, some were found (as will hereafter be more fully described) to be absolutely sterile when left to fertilise themselves spontaneously, although plenty of pollen fell on their stigmas; and they were equally sterile when artificially and repeatedly fertilised with their own pollen; whilst other plants produced a few spontaneously self-fertilised capsules. The remaining plants were left uncovered, and as pollen was carried from plant to plant by the hive and humble-bees which incessantly visit the flowers, they produced an abundance of capsules. Of the necessity of pollen being carried from one plant to another, I had ample evidence in the case of this species and of R. odorata; for those plants, which set no seeds or very few as long as they were protected from insects, became loaded with capsules immediately that they were uncovered.

Seeds from the flowers spontaneously self-fertilised under the net, and from flowers naturally crossed by the bees, were sown on opposite sides of five large pots. The seedlings were thinned as soon as they appeared above ground, so that an equal number were left on the two sides. After a time the pots were plunged into the open ground. The same number of plants of crossed and self-fertilised parentage were measured up to the summits of their flower-stems, with the result given in Table 4/35. Those which did not produce flower-stems were not measured.

TABLE 4/35. Reseda lutea, in pots.

Heights of plants to the summits of the flower-stems measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 21 : 12 7/8. Pot 1 : 14 2/8 : 16. Pot 1 : 19 1/8 : 11 7/8. Pot 1 : 7 : 15 2/8. Pot 1 : 15 1/8 : 19 1/8.

Pot 2 : 20 4/8 : 12 4/8. Pot 2 : 17 3/8 : 16 2/8. Pot 2 : 23 7/8 : 16 2/8. Pot 2 : 17 1/8 : 13 3/8. Pot 2 : 20 6/8 : 13 5/8.

Pot 3 : 16 1/8 : 14 4/8. Pot 3 : 17 6/8 : 19 4/8. Pot 3 : 16 2/8 : 20 7/8. Pot 3 : 10 : 7 7/8. Pot 3 : 10 : 17 6/8.

Pot 4 : 22 1/8 : 9. Pot 4 : 19 : 11 4/8. Pot 4 : 18 7/8 : 11. Pot 4 : 16 4/8 : 16. Pot 4 : 19 2/8 : 16 3/8.

Pot 5 : 25 2/8 : 14 6/8. Pot 5 : 22 : 16. Pot 5 : 8 6/8 : 14 3/8. Pot 5 : 14 2/8 : 14 2/8.

Total : 412.25 : 350.86.

The average height of the twenty-four crossed plants is here 17.17 inches, and that of the same number of self-fertilised plants 14.61; or as 100 to 85. Of the crossed plants all but five flowered, whilst several of the self-fertilised did not do so. The above pairs, whilst still in flower, but with some capsules already formed, were afterwards cut down and weighed. The crossed weighed 90.5 ounces; and an equal number of the self-fertilised only 19 ounces, or as 100 to 21; and this is an astonishing difference.

Seeds of the same two lots were also sown in two adjoining rows in the open ground. There were twenty crossed plants in the one row and thirty-two self-fertilised plants in the other row, so that the experiment was not quite fair; but not so unfair as it at first appears, for the plants in the same row were not crowded so much as seriously to interfere with each other’s growth, and the ground was bare on the outside of both rows. These plants were better nourished than those in the pots and grew to a greater height. The eight tallest plants in each row were measured in the same manner as before, with the following result:—

TABLE 4/36. Reseda lutea, growing in the open ground.

Heights of plants to the summits of the flower-stems measured in inches.

Column 1: Crossed Plants.

Column 2: Self-fertilised Plants.

28 : 33 2/8.
27 3/8 : 23.
27 5/8 : 21 5/8.
28 6/8 : 20 4/8.
29 7/8 : 21 5/8.
26 6/8 : 22.
26 2/8 : 21 2/8.
30 1/8 : 21 7/8.

Total : 224.75 : 185.13

The average height of the crossed plants, whilst in full flower, was here 28.09, and that of the self-fertilised 23.14 inches; or as 100 to 82. It is a singular fact that the tallest plant in the two rows, was one of the self-fertilised. The self-fertilised plants had smaller and paler green leaves than the crossed. All the plants in the two rows were afterwards cut down and weighed. The twenty crossed plants weighed 65 ounces, and twenty self-fertilised (by calculation from the actual weight of the thirty-two self-fertilised plants) weighed 26.25 ounces; or as 100 to 40. Therefore the crossed plants did not exceed in weight the self-fertilised plants in nearly so great a degree as those growing in the pots, owing probably to the latter having been subjected to more severe mutual competition. On the other hand, they exceeded the self-fertilised in height in a slightly greater degree.

Reseda odorata.

Plants of the common mignonette were raised from purchased seed, and several of them were placed under separate nets. Of these some became loaded with spontaneously self-fertilised capsules; others produced a few, and others not a single one. It must not be supposed that these latter plants produced no seed because their stigmas did not receive any pollen, for they were repeatedly fertilised with pollen from the same plant with no effect; but they were perfectly fertile with pollen from any other plant. Spontaneously self-fertilised seeds were saved from one of the highly self-fertile plants, and other seeds were collected from the plants growing outside the nets, which had been crossed by the bees. These seeds after germinating on sand were planted in pairs on the opposite sides of five pots. The plants were trained up sticks, and measured to the summits of their leafy stems—the flower-stems not being included. We here have the result:—

TABLE 4/37. Reseda odorata (seedlings from a highly self-fertile plant).

Heights of plants to the summits of the leafy stems, flower-stems not included, measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 20 7/8 : 22 4/8. Pot 1 : 34 7/8 : 28 5/8. Pot 1 : 26 6/8 : 23 2/8. Pot 1 : 32 6/8 : 30 4/8.

Pot 2 : 34 3/8 : 28 5/8. Pot 2 : 34 5/8 : 30 5/8. Pot 2 : 11 6/8 : 23. Pot 2 : 33 3/8 : 30 1/8.

Pot 3 : 17 7/8 : 4 4/8. Pot 3 : 27 : 25. Pot 3 : 30 1/8 : 26 3/8. Pot 3 : 30 2/8 : 25 1/8.

Pot 4 : 21 5/8 : 22 6/8. Pot 4 : 28 : 25 4/8. Pot 4 : 32 5/8 : 15 1/8. Pot 4 : 32 3/8 : 24 6/8.

Pot 5 : 21 : 11 6/8. Pot 5 : 25 2/8 : 19 7/8. Pot 5 : 26 6/8 : 10 4/8.

Total : 522.25 : 428.50.

The average height of the nineteen crossed plants is here 27.48, and that of the nineteen self-fertilised 22.55 inches; or as 100 to 82. All these plants were cut down in the early autumn and weighed: the crossed weighed 11.5 ounces, and the self-fertilised 7.75 ounces, or as 100 to 67. These two lots having been left freely exposed to the visits of insects, did not present any difference to the eye in the number of seed-capsules which they produced.

The remainder of the same two lots of seeds were sown in two adjoining rows in the open ground; so that the plants were exposed to only moderate competition. The eight tallest on each side were measured, as shown in Table 4/38.

TABLE 4/38. Reseda odorata, growing in the open ground.

Heights of plants measured in inches.

Column 1: Crossed Plants.

Column 2: Self-fertilised Plants.

24 4/8 : 26 5/8.
27 2/8 : 25 7/8.
24 : 25.
26 6/8 : 28 3/8.
25 : 29 7/8.
26 2/8 : 25 7/8.
27 2/8 : 26 7/8.
25 1/8 : 28 2/8.

Total : 206.13 : 216.75

The average height of the eight crossed plants is 25.76, and that of the eight self-fertilised 27.09; or as 100 to 105.

We here have the anomalous result of the self-fertilised plants being a little taller than the crossed; of which fact I can offer no explanation. It is of course possible, but not probable, that the labels may have been interchanged by accident.

Another experiment was now tried: all the self-fertilised capsules, though very few in number, were gathered from one of the semi-self-sterile plants under a net; and as several flowers on this same plant had been fertilised with pollen from a distinct individual, crossed seeds were thus obtained. I expected that the seedlings from this semi-self-sterile plant would have profited in a higher degree from a cross, than did the seedlings from the fully self-fertile plants. But my anticipation was quite wrong, for they profited in a less degree. An analogous result followed in the case of Eschscholtzia, in which the offspring of the plants of Brazilian parentage (which were partially self-sterile) did not profit more from a cross, than did the plants of the far more self-fertile English stock. The above two lots of crossed and self-fertilised seeds from the same plant of Reseda odorata, after germinating on sand, were planted on opposite sides of five pots, and measured as in the last case, with the result in Table 4/39.

TABLE 4/39. Reseda odorata (seedlings from a semi-self-sterile plant).

Heights of plants to the summits of the leafy stems, flower-stems not included, measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 33 4/8 : 31. Pot 1 : 30 6/8 : 28. Pot 1 : 29 6/8 : 13 2/8. Pot 1 : 20 : 32.

Pot 2 : 22 : 21 6/8. Pot 2 : 33 4/8 : 26 6/8. Pot 2 : 31 2/8 : 25 2/8. Pot 2 : 32 4/8 : 30 4/8.

Pot 3 : 30 1/8 : 17 2/8. Pot 3 : 32 1/8 : 29 6/8. Pot 3 : 31 4/8 : 24 6/8. Pot 3 : 32 2/8 : 34 2/8.

Pot 4 : 19 1/8 : 20 6/8. Pot 4 : 30 1/8 : 32 6/8. Pot 4 : 24 3/8 : 31 4/8. Pot 4 : 30 6/8 : 36 6/8.

Pot 5 : 34 6/8 : 24 5/8. Pot 5 : 37 1/8 : 34. Pot 5 : 31 2/8 : 22 2/8. Pot 5 : 33 : 37 1/8.

Total : 599.75 : 554.25.

The average height of the twenty crossed plants is here 29.98, and that of the twenty self-fertilised 27.71 inches; or as 100 to 92. These plants were then cut down and weighed; and the crossed in this case exceeded the self-fertilised in weight by a mere trifle, namely, in the ratio of 100 to 99. The two lots, left freely exposed to insects, seemed to be equally fertile.

The remainder of the seed was sown in two adjoining rows in the open ground; and the eight tallest plants in each row were measured, with the result in Table 4/40.

TABLE 4/40. Reseda odorata, (seedlings from a semi-self-sterile plant, planted in the open ground).

Heights of plants measured in inches.

Column 1: Crossed Plants.

Column 2: Self-fertilised Plants.

28 2/8 : 22 3/8.
22 4/8 : 24 3/8.
25 7/8 : 23 4/8.
25 3/8 : 21 4/8.
29 4/8 : 22 5/8.
27 1/8 : 27 3/8.
22 4/8 : 27 3/8.
26 2/8 : 19 2/8.

Total : 207.38 : 188.38.

The average height of the eight crossed plants is here 25.92, and that of the eight self-fertilised plants 23.54 inches; or as 100 to 90.

9. VIOLACEAE.—Viola tricolor.

Whilst the flowers of the common cultivated heartsease are young, the anthers shed their pollen into a little semi-cylindrical passage, formed by the basal portion of the lower petal, and surrounded by papillae. The pollen thus collected lies close beneath the stigma, but can seldom gain access into its cavity, except by the aid of insects, which pass their proboscides down this passage into the nectary. (4/5. The flowers of this plant have been fully described by Sprengel, Hildebrand, Delpino, and H. Muller. The latter author sums up all the previous observations in his ‘Befruchtung der Blumen’ and in ‘Nature’ November 20, 1873 page 44. See also Mr. A.W. Bennett in ‘Nature’ May 15, 1873 page 50 and some remarks by Mr. Kitchener ibid page 143. The facts which follow on the effects of covering up a plant of V. tricolor have been quoted by Sir J. Lubbock in his ‘British Wild Flowers’ etc. page 62.) Consequently when I covered up a large plant of a cultivated variety, it set only eighteen capsules, and most of these contained very few good seeds—several from only one to three; whereas an equally fine uncovered plant of the same variety, growing close by, produced 105 fine capsules. The few flowers which produce capsules when insects are excluded, are perhaps fertilised by the curling inwards of the petals as their wither, for by this means pollen-grains adhering to the papillae might be inserted into the cavity of the stigma. But it is more probable that their fertilisation is effected, as Mr. Bennett suggests, by Thrips and certain minute beetles which haunt the flowers, and which cannot be excluded by any net. Humble-bees are the usual fertilisers; but I have more than once seen flies (Rhingia rostrata) at work, with the under sides of their bodies, heads and legs dusted with pollen; and having marked the flowers which they visited, I found them after a few days fertilised. (4/6. I should add that this fly apparently did not suck the nectar, but was attracted by the papillae which surround the stigma. Hermann Muller also saw a small bee, an Andrena, which could not reach the nectar, repeatedly inserting its proboscis beneath the stigma, where the papillae are situated; so that these papillae must be in some way attractive to insects. A writer asserts ‘Zoologist’ volume 3-4 page 1225, that a moth (Plusia) frequently visits the flowers of the pansy. Hive-bees do not ordinarily visit them, but a case has been recorded ‘Gardeners’ Chronicle’ 1844 page 374, of these bees doing so. Hermann Muller has also seen the hive-bee at work, but only on the wild small-flowered form. He gives a list ‘Nature’ 1873 page 45, of all the insects which he has seen visiting both the large and small-flowered forms. From his account, I suspect that the flowers of plants in a state of nature are visited more frequently by insects than those of the cultivated varieties. He has seen several butterflies sucking the flowers of wild plants, and this I have never observed in gardens, though I have watched the flowers during many years.) It is curious for how long a time the flowers of the heartsease and of some other plants may be watched without an insect being seen to visit them. During the summer of 1841, I observed many times daily for more than a fortnight some large clumps of heartsease growing in my garden, before I saw a single humble-bee at work. During another summer I did the same, but at last saw some dark-coloured humble-bees visiting on three successive days almost every flower in several clumps; and almost all these flowers quickly withered and produced fine capsules. I presume that a certain state of the atmosphere is necessary for the secretion of nectar, and that as soon as this occurs the insects discover the fact by the odour emitted, and immediately frequent the flowers.

As the flowers require the aid of insects for their complete fertilisation, and as they are not visited by insects nearly so often as most other nectar-secreting flowers, we can understand the remarkable fact discovered by H. Muller and described by him in ‘Nature,’ namely, that this species exists under two forms. One of these bears conspicuous flowers, which, as we have seen, require the aid of insects, and are adapted to be cross-fertilised by them; whilst the other form has much smaller and less conspicuously coloured flowers, which are constructed on a slightly different plan, favouring self-fertilisation, and are thus adapted to ensure the propagation of the species. The self-fertile form, however, is occasionally visited, and may be crossed by insects, though this is rather doubtful.

In my first experiments on Viola tricolor I was unsuccessful in raising seedlings, and obtained only one full-grown crossed and self-fertilised plant. The former was 12 1/2 inches and the latter 8 inches in height. On the following year several flowers on a fresh plant were crossed with pollen from another plant, which was known to be a distinct seedling; and to this point it is important to attend. Several other flowers on the same plant were fertilised with their own pollen. The average number of seeds in the ten crossed capsules was 18.7, and in the twelve self-fertilised capsules 12.83; or as 100 to 69. These seeds, after germinating on bare sand, were planted in pairs on the opposite sides of five pots. They were first measured when about a third of their full size, and the crossed plants then averaged 3.87 inches, and the self-fertilised only 2.00 inches in height; or as 100 to 52. They were kept in the greenhouse, and did not grow vigorously. Whilst in flower they were again measured to the summits of their stems (see Table 4/41), with the following result:—

TABLE 4/41. Viola tricolor.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 8 2/8 : 0 2/8. Pot 1 : 7 4/8 : 2 4/8. Pot 1 : 5 : 1 2/8.

Pot 2 : 5 : 6. Pot 2 : 4 : 4. Pot 2 : 4 4/8 : 3 1/8.

Pot 3 : 9 4/8 : 3 1/8. Pot 3 : 3 3/8 : 1 7/8. Pot 3 : 8 4/8 : 0 5/8.

Pot 4 : 4 7/8 : 2 1/8. Pot 4 : 4 2/8 : 1 6/8. Pot 4 : 4 : 2 1/8.

Pot 5 : 6 : 3. Pot 5 : 3 3/8 : 1 4/8.

Total : 78.13 : 33.25.

The average height of the fourteen crossed plants is here 5.58 inches, and that of the fourteen self-fertilised 2.37; or as 100 to 42. In four out of the five pots, a crossed plant flowered before any one of the self-fertilised; as likewise occurred with the pair raised during the previous year. These plants without being disturbed were now turned out of their pots and planted in the open ground, so as to form five separate clumps. Early in the following summer (1869) they flowered profusely, and being visited by humble-bees set many capsules, which were carefully collected from all the plants on both sides. The crossed plants produced 167 capsules, and the self-fertilised only 17; or as 100 to 10. So that the crossed plants were more than twice the height of the self-fertilised, generally flowered first, and produced ten times as many naturally fertilised capsules.

By the early part of the summer of 1870 the crossed plants in all the five clumps had grown and spread so much more than the self-fertilised, that any comparison between them was superfluous. The crossed plants were covered with a sheet of bloom, whilst only a single self-fertilised plant, which was much finer than any of its brethren, flowered. The crossed and self-fertilised plants had now grown all matted together on the respective sides of the superficial partitions still separating them; and in the clump which included the finest self-fertilised plant, I estimated that the surface covered by the crossed plants was about nine times as large as that covered by the self-fertilised plants. The extraordinary superiority of the crossed over the self-fertilised plants in all five clumps, was no doubt due to the crossed plants at first having had a decided advantage over the self-fertilised, and then robbing them more and more of their food during the succeeding seasons. But we should remember that the same result would follow in a state of nature even to a greater degree; for my plants grew in ground kept clear of weeds, so that the self-fertilised had to compete only with the crossed plants; whereas the whole surface of the ground is naturally covered with various kinds of plants, all of which have to struggle together for existence.

The ensuing winter was very severe, and in the following spring (1871) the plants were again examined. All the self-fertilised were now dead, with the exception of a single branch on one plant, which bore on its summit a minute rosette of leaves about as large as a pea. On the other hand, all the crossed plants without exception were growing vigorously. So that the self-fertilised plants, besides their inferiority in other respects, were more tender.

Another experiment was now tried for the sake of ascertaining how far the superiority of the crossed plants, or to speak more correctly, the inferiority of the self-fertilised plants, would be transmitted to their offspring. The one crossed and one self-fertilised plant, which were first raised, had been turned out of their pot and planted in the open ground. Both produced an abundance of very fine capsules, from which fact we may safely conclude that they had been cross-fertilised by insects. Seeds from both, after germinating on sand, were planted in pairs on the opposite sides of three pots. The naturally crossed seedlings derived from the crossed plants flowered in all three pots before the naturally crossed seedlings derived from the self-fertilised plants. When both lots were in full flower, the two tallest plants on each side of each pot were measured, and the result is shown in Table 4/42.

TABLE 4/42. Viola tricolor: seedlings from crossed and self-fertilised plants, the parents of both sets having been left to be naturally fertilised.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Naturally Crossed Plants from artificially crossed Plants.

Column 3: Naturally Crossed Plants from Self-fertilised Plants.

Pot 1 : 12 1/8 : 9 6/8. Pot 1 : 11 6/8 : 8 3/8.

Pot 2 : 13 2/8 : 9 6/8. Pot 2 : 10 : 11 4/8.

Pot 3 : 14 4/8 : 11 1/8. Pot 3 : 13 6/8 : 11 3/8.

Total : 75.38 : 61.88.

The average height of the six tallest plants derived from the crossed plants is 12.56 inches; and that of the six tallest plants derived from the self-fertilised plants is 10.31 inches; or as 100 to 82. We here see a considerable difference in height between the two sets, though very far from equalling that in the previous trials between the offspring from crossed and self-fertilised flowers. This difference must be attributed to the latter set of plants having inherited a weak constitution from their parents, the offspring of self-fertilised flowers; notwithstanding that the parents themselves had been freely intercrossed with other plants by the aid of insects.

10. RANUNCULACEAE.—Adonis aestivalis.

The results of my experiments on this plant are hardly worth giving, as I remark in my notes made at the time, “seedlings, from some unknown cause, all miserably unhealthy.” Nor did they ever become healthy; yet I feel bound to give the present case, as it is opposed to the general results at which I have arrived. Fifteen flowers were crossed and all produced fruit, containing on an average 32.5 seeds; nineteen flowers were fertilised with their own pollen, and they likewise all yielded fruit, containing a rather larger average of 34.5 seeds; or as 100 to 106. Seedlings were raised from these seeds. In one of the pots all the self-fertilised plants died whilst quite young; in the two others, the measurements were as follows:

TABLE 4/43. Adonis aestivalis.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 14 : 13 4/8. Pot 1 : 13 4/8 : 13 4/8.

Pot 2 : 16 2/8 : 15 2/8. Pot 2 : 13 2/8 : 15.

Total : 57.00 : 57.25.

The average height of the four crossed plants is 14.25, and that of the four self-fertilised plants 14.31; or as 100 to 100.4; so that they were in fact of equal height. According to Professor H. Hoffman, this plant is proterandrous (4/7. ‘Zur Speciesfrage’ 1875 page 11.); nevertheless it yields plenty of seeds when protected from insects.

Delphinium consolida.

It has been said in the case of this plant, as of so many others, that the flowers are fertilised in the bud, and that distinct plants or varieties can never naturally intercross. (4/8. Decaisne ‘Comptes-Rendus’ July 1863 page 5.) But this is an error, as we may infer, firstly from the flowers being proterandrous,—the mature stamens bending up, one after the other, into the passage which leads to the nectary, and afterwards the mature pistils bending in the same direction; secondly, from the number of humble-bees which visit the flowers (4/9. Their structure is described by H. Muller ‘Befruchtung’ etc., page 122.); and thirdly, from the greater fertility of the flowers when crossed with pollen from a distinct plant than when spontaneously self-fertilised. In the year 1863 I enclosed a large branch in a net, and crossed five flowers with pollen from a distinct plant; these yielded capsules containing on an average 35.2 very fine seeds, with a maximum of forty-two in one capsule. Thirty-two other flowers on the same branch produced twenty-eight spontaneously self-fertilised capsules, containing on an average 17.2 seeds, with a maximum in one of thirty-six seeds. But six of these capsules were very poor, yielding only from one to five seeds; if these are excluded, the remaining twenty-two capsules give an average of 20.9 seeds, though many of these seeds were small. The fairest ratio, therefore, for the number of seeds produced by a cross and by spontaneous self-fertilisation is as 100 to 59. These seeds were not sown, as I had too many other experiments in progress.

In the summer of 1867, which was a very unfavourable one, I again crossed several flowers under a net with pollen from a distinct plant, and fertilised other flowers on the same plant with their own pollen. The former yielded a much larger proportion of capsules than the latter; and many of the seeds in the self-fertilised capsules, though numerous, were so poor that an equal number of seeds from the crossed and self-fertilised capsules were in weight as 100 to 45. The two lots were allowed to germinate on sand, and pairs were planted on the opposite sides of four pots. When nearly two-thirds grown they were measured, as shown in Table 4/44.

TABLE 4/44. Delphinium consolida.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 11 : 11.

Pot 2 : 19 : 16 2/8. Pot 2 : 16 2/8 : 11 4/8.

Pot 3 : 26 : 22.

Pot 4 : 9 4/8 : 8 2/8. Pot 4 : 8 : 6 4/8.

Total : 89.75 : 75.50.

The six crossed plants here average 14.95, and the six self-fertilised 12.50 inches in height; or as 100 to 84. When fully grown they were again measured, but from want of time only a single plant on each side was measured; so that I have thought it best to give the earlier measurements. At the later period the three tallest crossed plants still exceeded considerably in height the three tallest self-fertilised, but not in quite so great a degree as before. The pots were left uncovered in the greenhouse, but whether the flowers were intercrossed by bees or self-fertilised I do not know. The six crossed plants produced 282 mature and immature capsules, whilst the six self-fertilised plants produced only 159; or as 100 to 56. So that the crossed plants were very much more productive than the self-fertilised.

11. CARYOPHYLLACEAE.—Viscaria oculata.

Twelve flowers were crossed with pollen from another plant, and yielded ten capsules, containing by weight 5.77 grains of seeds. Eighteen flowers were fertilised with their own pollen and yielded twelve capsules, containing by weight 2.63 grains. Therefore the seeds from an equal number of crossed and self-fertilised flowers would have been in weight as 100 to 38. I had previously selected a medium-sized capsule from each lot, and counted the seeds in both; the crossed one contained 284, and the self-fertilised one 126 seeds; or as 100 to 44. These seeds were sown on opposite sides of three pots, and several seedlings raised; but only the tallest flower-stem of one plant on each side was measured. The three on the crossed side averaged 32.5 inches, and the three on the self-fertilised side 34 inches in height; or as 100 to 104. But this trial was on much too small a scale to be trusted; the plants also grew so unequally that one of the three flower-stems on the crossed plants was very nearly twice as tall as that on one of the others; and one of the three flower-stems on the self-fertilised plants exceeded in an equal degree one of the others.

In the following year the experiment was repeated on a larger scale: ten flowers were crossed on a new set of plants and yielded ten capsules containing by weight 6.54 grains of seed. Eighteen spontaneously self-fertilised capsules were gathered, of which two contained no seed; the other sixteen contained by weight 6.07 grains of seed. Therefore the weight of seed from an equal number of crossed and spontaneously self-fertilised flowers (instead of artificially fertilised as in the previous case) was as 100 to 58.

The seeds after germinating on sand were planted in pairs on the opposite sides of four pots, with all the remaining seeds sown crowded in the opposite sides of a fifth pot; in this latter pot only the tallest plant on each side was measured. Until the seedlings had grown about 5 inches in height no difference could be perceived in the two lots. Both lots flowered at nearly the same time. When they had almost done flowering, the tallest flower-stem on each plant was measured, as shown in Table 4/45.

TABLE 4/45. Viscaria oculata.

Tallest flower-stem on each plant measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 19 : 32 3/8. Pot 1 : 33 : 38. Pot 1 : 41 : 38. Pot 1 : 41 : 28 7/8.

Pot 2 : 37 4/8 : 36. Pot 2 : 36 4/8 : 32 3/8. Pot 2 : 38 : 35 6/8.

Pot 3 : 44 4/8 : 36. Pot 3 : 39 4/8 : 20 7/8. Pot 3 : 39 : 30 5/8.

Pot 4 : 30 2/8 : 36. Pot 4 : 31 : 39. Pot 4 : 33 1/8 : 29. Pot 4 : 24 : 38 4/8.

Pot 5 : 30 2/8 : 32. Crowded.

Total : 517.63 : 503.36.

The fifteen crossed plants here average 34.5, and the fifteen self-fertilised 33.55 inches in height; or as 100 to 97. So that the excess of height of the crossed plants is quite insignificant. In productiveness, however, the difference was much more plainly marked. All the capsules were gathered from both lots of plants (except from the crowded and unproductive ones in Pot 5), and at the close of the season the few remaining flowers were added in. The fourteen crossed plants produced 381, whilst the fourteen self-fertilised plants produced only 293 capsules and flowers; or as 100 to 77.

Dianthus caryophyllus.

The common carnation is strongly proterandrous, and therefore depends to a large extent upon insects for fertilisation. I have seen only humble-bees visiting the flowers, but I dare say other insects likewise do so. It is notorious that if pure seed is desired, the greatest care is necessary to prevent the varieties which grow in the same garden from intercrossing. (4/10. ‘Gardeners’ Chronicle’ 1847 page 268.) The pollen is generally shed and lost before the two stigmas in the same flower diverge and are ready to be fertilised. I was therefore often forced to use for self-fertilisation pollen from the same plant instead of from the same flower. But on two occasions, when I attended to this point, I was not able to detect any marked difference in the number of seeds produced by these two forms of self-fertilisation.

Several single-flowered carnations were planted in good soil, and were all covered with a net. Eight flowers were crossed with pollen from a distinct plant and yielded six capsules, containing on an average 88.6 seeds, with a maximum in one of 112 seeds. Eight other flowers were self-fertilised in the manner above described, and yielded seven capsules containing on an average 82 seeds, with a maximum in one of 112 seeds. So that there was very little difference in the number of seeds produced by cross-fertilisation and self-fertilisation, namely, as 100 to 92. As these plants were covered by a net, they produced spontaneously only a few capsules containing any seeds, and these few may perhaps be attributed to the action of Thrips and other minute insects which haunt the flowers. A large majority of the spontaneously self-fertilised capsules produced by several plants contained no seeds, or only a single one. Excluding these latter capsules, I counted the seeds in eighteen of the finest ones, and these contained on an average 18 seeds. One of the plants was spontaneously self-fertile in a higher degree than any of the others. On another occasion a single covered-up plant produced spontaneously eighteen capsules, but only two of these contained any seed, namely 10 and 15.

CROSSED AND SELF-FERTILISED PLANTS OF THE FIRST GENERATION.

The many seeds obtained from the above crossed and artificially self-fertilised flowers were sown out of doors, and two large beds of seedlings, closely adjoining one another, thus raised. This was the first plant on which I experimented, and I had not then formed any regular scheme of operation. When the two lots were in full flower, I measured roughly a large number of plants but record only that the crossed were on an average fully 4 inches taller than the self-fertilised. Judging from subsequent measurements, we may assume that the crossed plants were about 28 inches, and the self-fertilised about 24 inches in height; and this will give us a ratio of 100 to 86. Out of a large number of plants, four of the crossed ones flowered before any one of the self-fertilised plants.

Thirty flowers on these crossed plants of the first generation were again crossed with pollen from a distinct plant of the same lot, and yielded twenty-nine capsules, containing on an average 55.62 seeds, with a maximum in one of 110 seeds.

Thirty flowers on the self-fertilised plants were again self-fertilised; eight of them with pollen from the same flower, and the remainder with pollen from another flower on the same plant; and these produced twenty-two capsules, containing on an average 35.95 seeds, with a maximum in one of sixty-one seeds. We thus see, judging by the number of seeds per capsule, that the crossed plants again crossed were more productive than the self-fertilised again self-fertilised, in the ratio of 100 to 65. Both the crossed and self-fertilised plants, from having grown much crowded in the two beds, produced less fine capsules and fewer seeds than did their parents.

CROSSED AND SELF-FERTILISED PLANTS OF THE SECOND GENERATION.

The crossed and self-fertilised seeds from the crossed and self-fertilised plants of the last generation were sown on opposite sides of two pots; but the seedlings were not thinned enough, so that both lots grew very irregularly, and most of the self-fertilised plants after a time died from being smothered. My measurements were, therefore, very incomplete. From the first the crossed seedlings appeared the finest, and when they were on an average, by estimation, 5 inches high, the self-fertilised plants were only 4 inches. In both pots the crossed plants flowered first. The two tallest flower-stems on the crossed plants in the two pots were 17 and 16 1/2 inches in height; and the two tallest flower-stems on the self-fertilised plants 10 1/2 and 9 inches; so that their heights were as 100 to 58. But this ratio, deduced from only two pairs, obviously is not in the least trustworthy, and would not have been given had it not been otherwise supported. I state in my notes that the crossed plants were very much more luxuriant than their opponents, and seemed to be twice as bulky. This latter estimate may be believed from the ascertained weights of the two lots in the next generation. Some flowers on these crossed plants were again crossed with pollen from another plant of the same lot, and some flowers on the self-fertilised plants again self-fertilised; and from the seeds thus obtained the plants of the next generation were raised.

CROSSED AND SELF-FERTILISED PLANTS OF THE THIRD GENERATION.

The seeds just alluded to were allowed to germinate on bare sand, and were planted in pairs on the opposite sides of four pots. When the seedlings were in full flower, the tallest stem on each plant was measured to the base of the calyx. The measurements are given in Table 4/46. In Pot 1 the crossed and self-fertilised plants flowered at the same time; but in the other three pots the crossed flowered first. These latter plants also continued flowering much later in the autumn than the self-fertilised.

TABLE 4/46. Dianthus caryophyllus (third generation).

Tallest flower-stem on each plant measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 28 6/8 : 30. Pot 1 : 27 3/8 : 26.

Pot 2 : 29 : 30 7/8. Pot 2 : 29 4/8 : 27 4/8.

Pot 3 : 28 4/8 : 31 6/8. Pot 3 : 23 4/8 : 24 5/8.

Pot 4 : 27 : 30. Pot 4 : 33 4/8 : 25.

Total : 227.13 : 225.75.

The average height of the eight crossed plants is here 28.39 inches, and of the eight self-fertilised 28.21; or as 100 to 99. So that there was no difference in height worth speaking of; but in general vigour and luxuriance there was an astonishing difference, as shown by their weights. After the seed-capsules had been gathered, the eight crossed and the eight self-fertilised plants were cut down and weighed; the former weighed 43 ounces, and the latter only 21 ounces; or as 100 to 49.

These plants were all kept under a net, so that the capsules which they produced must have been all spontaneously self-fertilised. The eight crossed plants produced twenty-one such capsules, of which only twelve contained any seed, averaging 8.5 per capsule. On the other hand, the eight self-fertilised plants produced no less than thirty-six capsules, of which I examined twenty-five, and, with the exception of three, all contained seeds, averaging 10.63 seeds per capsule. Thus the proportional number of seeds per capsule produced by the plants of crossed origin to those produced by the plants of self-fertilised origin (both lots being spontaneously self-fertilised) was as 100 to 125. This anomalous result is probably due to some of the self-fertilised plants having varied so as to mature their pollen and stigmas more nearly at the same time than is proper to the species; and we have already seen that some plants in the first experiment differed from the others in being slightly more self-fertile.

THE EFFECTS OF A CROSS WITH A FRESH STOCK.

Twenty flowers on the self-fertilised plants of the last or third generation, in Table 4/46, were fertilised with their own pollen, but taken from other flowers on the same plants. These produced fifteen capsules, which contained (omitting two with only three and six seeds) on an average 47.23 seeds, with a maximum of seventy in one. The self-fertilised capsules from the self-fertilised plants of the first generation yielded the much lower average of 35.95 seeds; but as these latter plants grew extremely crowded, nothing can be inferred with respect to this difference in their self-fertility. The seedlings raised from the above seeds constitute the plants of the fourth self-fertilised generation in Table 4/47.

Twelve flowers on the same plants of the third self-fertilised generation, in Table 4/46, were crossed with pollen from the crossed plants in the same table. These crossed plants had been intercrossed for the three previous generations; and many of them, no doubt, were more or less closely inter-related, but not so closely as in some of the experiments with other species; for several carnation plants had been raised and crossed in the earlier generations. They were not related, or only in a distant degree, to the self-fertilised plants. The parents of both the self-fertilised and crossed plants had been subjected to as nearly as possible the same conditions during the three previous generations. The above twelve flowers produced ten capsules, containing on an average 48.66 seeds, with a maximum in one of seventy-two seeds. The plants raised from these seeds may be called the INTERCROSSED.

Lastly, twelve flowers on the same self-fertilised plants of the third generation were crossed with pollen from plants which had been raised from seeds purchased in London. It is almost certain that the plants which produced these seeds had grown under very different conditions to those to which my self-fertilised and crossed plants had been subjected; and they were in no degree related. The above twelve flowers thus crossed all produced capsules, but these contained the low average of 37.41 seeds per capsule, with a maximum in one of sixty-four seeds. It is surprising that this cross with a fresh stock did not give a much higher average number of seeds; for, as we shall immediately see, the plants raised from these seeds, which may be called the LONDON-CROSSED, benefited greatly by the cross, both in growth and fertility.

The above three lots of seeds were allowed to germinate on bare sand. Many of the London-crossed germinated before the others, and were rejected; and many of the intercrossed later than those of the other two lots. The seeds after thus germinating were planted in ten pots, made tripartite by superficial divisions; but when only two kinds of seeds germinated at the same time, they were planted on the opposite sides of other pots; and this is indicated by blank spaces in one of the three columns in Table 4/47. A 0 in the table signifies that the seedling died before it was measured; and a + signifies that the plant did not produce a flower-stem, and therefore was not measured. It deserves notice that no less than eight out of the eighteen self-fertilised plants either died or did not flower; whereas only three out of the eighteen intercrossed, and four out of the twenty London-crossed plants, were in this predicament. The self-fertilised plants had a decidedly less vigorous appearance than the plants of the other two lots, their leaves being smaller and narrower. In only one pot did a self-fertilised plant flower before one of the two kinds of crossed plants, between which there was no marked difference in the period of flowering. The plants were measured to the base of the calyx, after they had completed their growth, late in the autumn.

TABLE 4/47. Dianthus caryophyllus.

Heights of plants to the base of the calyx, measured in inches.

Column 1: Number (Name) of Pot.

Column 2: London-Crossed Plants.

Column 3: Intercrossed Plants.

Column 4: Self-fertilised Plants.

Pot 1 : 39 5/8 : 25 1/8 : 29 2/8. Pot 1 : 30 7/8 : 21 6/8 : +.

Pot 2 : 36 2/8 : : 22 3/8. Pot 2 : 0 : : +.

Pot 3 : 28 5/8 : 30 2/8 : . Pot 3 : + : 23 1/8 : .

Pot 4 : 33 4/8 : 35 5/8 : 30. Pot 4 : 28 7/8 : 32 : 24 4/8.

Pot 5 : 28 : 34 4/8 : +. Pot 5 : 0 : 24 2/8 : +.

Pot 6 : 32 5/8 : 24 7/8 : 30 3/8. Pot 6 : 31 : 26 : 24 4/8.

Pot 7 : 41 7/8 : 29 7/8 : 27 7/8. Pot 7 : 34 7/8 : 26 4/8 : 27.

Pot 8 : 34 5/8 : 29 : 26 6/8. Pot 8 : 28 5/8 : 0 : +.

Pot 9 : 25 5/8 : 28 5/8 : +. Pot 9 : 0 : + : 0.

Pot 10 : 38 : 28 4/8 : 22 7/8. Pot 10 : 32 1/8 : + : 0.

Total : 525.13 : 420.00 : 265.50.

The average height of the sixteen London-crossed plants in Table 4/47 is 32.82 inches; that of the fifteen intercrossed plants, 28 inches; and that of the ten self-fertilised plants, 26.55.

So that in height we have the following ratios:—

The London-crossed to the self-fertilised as 100 to 81.

The London-crossed to the intercrossed as 100 to 85.

The intercrossed to the self-fertilised as 100 to 95.

These three lots of plants, which it should be remembered were all derived on the mother-side from plants of the third self-fertilised generation, fertilised in three different ways, were left exposed to the visits of insects, and their flowers were freely crossed by them. As the capsules of each lot became ripe they were gathered and kept separate, the empty or bad ones being thrown away. But towards the middle of October, when the capsules could no longer ripen, all were gathered and were counted, whether good or bad. The capsules were then crushed, and the seed cleaned by sieves and weighed. For the sake of uniformity the results are given from calculation, as if there had been twenty plants in each lot.

The sixteen London-crossed plants actually produced 286 capsules; therefore twenty such plants would have produced 357.5 capsules; and from the actual weight of the seeds, the twenty plants would have yielded 462 grains weight of seeds.

The fifteen intercrossed plants actually produced 157 capsules; therefore twenty of them would have produced 209.3 capsules and the seeds would have weighed 208.48 grains.

The ten self-fertilised plants actually produced 70 capsules, therefore twenty of them would have produced 140 capsules; and the seeds would have weighed 153.2 grains.

From these data we get the following ratios:—

NUMBER OF CAPSULES PRODUCED BY AN EQUAL NUMBER OF PLANTS OF THE THREE LOTS.

NUMBER OF CAPSULES:

The London-crossed to the self-fertilised as 100 to 39.

The London-crossed to the intercrossed as 100 to 45.

The intercrossed to the self-fertilised as 100 to 67.

WEIGHT OF SEEDS PRODUCED BY AN EQUAL NUMBER OF PLANTS OF THE THREE LOTS.

WEIGHT OF SEED:

The London-crossed to the self-fertilised as 100 to 33.

The London-crossed to the intercrossed as 100 to 45.

The intercrossed to the self-fertilised as 100 to 73.

We thus see how greatly the offspring from the self-fertilised plants of the third generation crossed by a fresh stock, had their fertility increased, whether tested by the number of capsules produced or by the weight of the contained seeds; this latter being the more trustworthy method. Even the offspring from the self-fertilised plants crossed by one of the crossed plants of the same stock, notwithstanding that both lots had been long subjected to the same conditions, had their fertility considerably increased, as tested by the same two methods.

In conclusion it may be well to repeat in reference to the fertility of these three lots of plants, that their flowers were left freely exposed to the visits of insects and were undoubtedly crossed by them, as may be inferred from the large number of good capsules produced. These plants were all the offspring of the same mother-plants, and the strongly marked difference in their fertility must be attributed to the nature of the pollen employed in fertilising their parents; and the difference in the nature of the pollen must be attributed to the different treatment to which the pollen-bearing parents had been subjected during several previous generations.

COLOUR OF THE FLOWERS.

The flowers produced by the self-fertilised plants of the last or fourth generation were as uniform in tint as those of a wild species, being of a pale pink or rose colour. Analogous cases with Mimulus and Ipomoea, after several generations of self-fertilisation, have been already given. The flowers of the intercrossed plants of the fourth generation were likewise nearly uniform in colour. On the other hand, the flowers of the London-crossed plants, or those raised from a cross with the fresh stock which bore dark crimson flowers, varied extremely in colour, as might have been expected, and as is the general rule with seedling carnations. It deserves notice that only two or three of the London-crossed plants produced dark crimson flowers like those of their fathers, and only a very few of a pale pink like those of their mothers. The great majority had their petals longitudinally and variously striped with the two colours,—the groundwork tint being, however, in some cases darker than that of the mother-plants.

12. MALVACEAE.—Hibiscus africanus.

Many flowers on this Hibiscus were crossed with pollen from a distinct plant, and many others were self-fertilised. A rather larger proportional number of the crossed than of the self-fertilised flowers yielded capsules, and the crossed capsules contained rather more seeds. The self-fertilised seeds were a little heavier than an equal number of the crossed seeds, but they germinated badly, and I raised only four plants of each lot. In three out of the four pots, the crossed plants flowered first.

TABLE 4/48. Hibiscus africanus.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 13 4/8 : 16 2/8.

Pot 2 : 14 : 14.

Pot 3 : 8 : 7.

Pot 4 : 17 4/8 : 20 4/8.

Total : 53.00 : 57.75.

The four crossed plants average 13.25, and the four self-fertilised 14.43 inches in height; or as 100 to 109. Here we have the unusual case of self-fertilised plants exceeding the crossed in height; but only four pairs were measured, and these did not grow well or equally. I did not compare the fertility of the two lots.

CHAPTER V. GERANIACEAE, LEGUMINOSAE, ONAGRACEAE, ETC.

Pelargonium zonale, a cross between plants propagated by cuttings does
no good.
Tropaeolum minus.
Limnanthes douglasii.
Lupinus luteus and pilosus.
Phaseolus multiflorus and vulgaris.
Lathyrus odoratus, varieties of, never naturally intercross in England.
Pisum sativum, varieties of, rarely intercross, but a cross between them
highly beneficial.
Sarothamnus scoparius, wonderful effects of a cross.
Ononis minutissima, cleistogene flowers of.
Summary on the Leguminosae.
Clarkia elegans.
Bartonia aurea.
Passiflora gracilis.
Apium petroselinum.
Scabiosa atropurpurea.
Lactuca sativa.
Specularia speculum.
Lobelia ramosa, advantages of a cross during two generations.
Lobelia fulgens.
Nemophila insignis, great advantages of a cross.
Borago officinalis.
Nolana prostrata.

13. GERANIACEAE.—Pelargonium zonale.

This plant, as a general rule, is strongly proterandrous, and is therefore adapted for cross-fertilisation by the aid of insects. (5/1. Mr. J. Denny, a great raiser of new varieties of pelargoniums, after stating that this species is proterandrous, adds ‘The Florist and Pomologist’ January 1872 page 11, “there are some varieties, especially those with petals of a pink colour, or which possess a weakly constitution, where the pistil expands as soon as or even before the pollen-bag bursts, and in which also the pistil is frequently short, so when it expands it is smothered as it were by the bursting anthers; these varieties are great seeders, each pip being fertilised by its own pollen. I would instance Christine as an example of this fact.” We have here an interesting case of variability in an important functional point.) Some flowers on a common scarlet variety were self-fertilised, and other flowers were crossed with pollen from another plant; but no sooner had I done so, than I remembered that these plants had been propagated by cuttings from the same stock, and were therefore parts in a strict sense of the same individual. Nevertheless, having made the cross I resolved to save the seeds, which, after germinating on sand, were planted on the opposite sides of three pots. In one pot the quasi-crossed plant was very soon and ever afterwards taller and finer than the self-fertilised. In the two other pots the seedlings on both sides were for a time exactly equal; but when the self-fertilised plants were about 10 inches in height, they surpassed their antagonists by a little, and ever afterwards showed a more decided and increasing advantage; so that the self-fertilised plants, taken altogether, were somewhat superior to the quasi-crossed plants. In this case, as in that of the Origanum, if individuals which have been asexually propagated from the same stock, and which have been long subjected to the same conditions, are crossed, no advantage whatever is gained.

Several flowers on another plant of the same variety were fertilised with pollen from the younger flowers on the same plant, so as to avoid using the old and long-shed pollen from the same flower, as I thought that this latter might be less efficient than fresh pollen. Other flowers on the same plant were crossed with fresh pollen from a plant which, although closely similar, was known to have arisen as a distinct seedling. The self-fertilised seeds germinated rather before the others; but as soon as I got equal pairs they were planted on the opposite sides of four pots.

TABLE 5/49. Pelargonium zonale.

Heights of plants measured in inches.

Column 1: Number (Name) of Pot.

Column 2: Crossed Plants.

Column 3: Self-fertilised Plants.

Pot 1 : 22 3/8 : 25 5/8. Pot 1 : 19 6/8 : 12 4/8.

Pot 2 : 15 : 19 6/8. Pot 2 : 12 2/8 : 22 3/8.

Pot 3 : 30 5/8 : 19 4/8. Pot 3 : 18 4/8 : 7 4/8.

Pot 4 : 38 : 9 1/8.

Total : 156.50 : 116.38.

When the two lots of seedlings were between 4 and 5 inches in height they were equal, excepting in Pot 4, in which the crossed plant was much the tallest. When between 11 and 14 inches in height, they were measured to the tips of their uppermost leaves; the crossed averaged 13.46, and the self-fertilised 11.07 inches in height, or as 100 to 82. Five months later they were again measured in the same manner, and the results are given in Table 5/49.

The seven crossed plants now averaged 22.35, and the seven self-fertilised 16.62 inches in height, or as 100 to 74. But from the great inequality of the several plants, the result is less trustworthy than in most other cases. In Pot 2 the two self-fertilised plants always had an advantage, except whilst quite young over the two crossed plants.