Personal Characteristics.
In as much as it is our custom to erect none but the slightest and most casual memorials to our distinguished men of science or of letters, there is reason to rejoice that the name of Lindley is not inadequately commemorated.
The Lindley Library purchased in his honour and now permanently attached to the Royal Horticultural Society bids fair under the enlightened policy of that flourishing institution to grow into a great collection of horticultural works. The genus Lindleya is reminiscent to systematists of their great colleague and the name of Lindley is known and honoured by all our horticulturists. Of the man himself just so much may be said as to give form to the mind's image of him.
He was of middle height, active, upright, with shoulders somewhat sloping and of heavy tread. The sightlessness of one eye gave to his resolute face a somewhat strange look. Simple in habits, strenuous in work and perspicacious in judgment, John Lindley was a warm hearted and generous friend, particularly to young botanists. He was a powerful foe: altogether a masterful and remarkable man. Not suffering fools gladly yet with a humorous turn of mind: "I am a dandy in my herbarium," he once exclaimed to Reichenbach. Knowing no fear he could not hope for much favour, and yet carrying his heavy load of financial responsibility, he nevertheless won through to a wide measure of contemporary recognition and an assured place in the history of botanical science. To conclude with Reichenbach's fine tribute "we cannot tell how long Botany, how long science, will be pursued; but we may affirm that so long as a knowledge of plants is considered necessary, so long will Lindley's name be remembered with gratitude."
[WILLIAM GRIFFITH]
1810-1845
By W. H. LANG
Early training—medical appointment under the East India Company—his travels—the magnitude of his collections—his method of work—results of researches mainly published posthumously—the ovule and fertilisation—Santalum—Loranthaceae—Balanophora—Avicennia—his gymnosperm work illustrated by Cycas—discovery of the pollen-chamber—Rhizocarps and Liverworts—pre-Hofmeisterian work—Griffith's relation to his times.
It might have been assumed that all the names of British botanists whose work has been or is to be considered in this course of lectures would have been familiar to their successors of to-day, even if their works were too often neglected for the last words of scientific progress in a summary of literature. The question has however been put to me by more than one botanist in the last month or two, "But who was Griffith?" That this should be possible seems in itself ample justification for including his name in this list of British botanists.
For Griffith has claims to be regarded as a great botanist. It is true that he failed to break through the limitations of his time and period—that he left no new and more correct general views to modify the science. But this is true of all his contemporaries, indeed it is true of most botanists. To recreate the department of a science in which a man labours requires a combination of ability and fortunate chance that is given to few.
Plate XV
WILLIAM GRIFFITH (1843)
Griffith had the ability, the power of independent observation, the readiness to speculate, the careless prodigality of labour. He did not however, in the fraction of an ordinary working life that fate allowed him, attain that insight into more correct comparison of the plants whose morphology he studied which would have acted quickly on the mass of first hand observation he possessed.
It is well to be clear at the outset that it is the personality of William Griffith, his important detailed contributions to botany, and his achievement as a great working morphologist of his time that will interest us to-day—rather than his general views or any influence of these on the progress of botany. Griffith had the advantage or disadvantage of botany being his private study and not his profession. The motive force of his career was however his love of scientific work for its own sake.
William Griffith was a London botanist. He was the son of a London merchant, born on March 4, 1810, at Ham Common. Having finished school he began to prepare for the medical profession and was apprenticed to a surgeon in the West end of London. About 1829 he commenced attendance at the classes in the newly established University College. He had earlier in life shown an interest in natural history but was now specially devoted to botany. He attended Lindley's lectures, and also studied medical botany under Mr Anderson at the Apothecaries' Garden in Chelsea. There he obtained the Linnean Gold Medal given by the Society of Apothecaries. At this time also he was a frequent visitor to Kew Gardens where he was on good terms with the head gardener and also came under the influence of Mr Bauer the great botanical draughtsman of his day. Griffith was never tired of expressing his admiration for Bauer as an accurate observer. During his vacations Griffith made botanical excursions in England, carrying his light baggage and his equipment for collecting plants.
That the training that Griffith received in botany in the London University of that date was a sound one is shown by his power of facing the most various problems when cast on his own resources immediately at the close of his University training. The soundness of his training is further shown by the small pieces of original work he had published before leaving England at the age of 22. Not only had he made some of the illustrations for Lindley's Introduction to Botany and had described the flower and the structure of the wood of Phytocrene gigantea in Wallich's Plantae Asiaticae Rariores, but (a noteworthy indication of his interest in Cryptogams at this time) he had supplied an account of the structure and development of Targionia hypophylla to be appended to Mirbel's classic monograph on the anatomy and physiology of Marchantia polymorpha—published in 1832.
His medical studies finished, Griffith sailed from England in May 1832, he arrived at Madras in September and was appointed Assistant-Surgeon on the Madras establishment in the service of the East India Company. His scientific work was done in the intervals of a busy life. Only a man of great energy and enthusiasm and possessed of great powers of physical endurance could have done the work that Griffith crowded into the 12½ years, between his landing in India and his death at Malacca before the age of 35 on February 9, 1845. This time was all spent in the East Indies—he never returned to England.
Deferring for the moment consideration of his scientific work we may take a general survey of Griffith's movements during his working life and of his labours as an explorer and collector.
After spending some months in the neighbourhood of Madras, he was situated for more than two years at Mergui and collected extensively in Tenasserim. He was recalled to Calcutta in 1835 and attached to the Bengal Presidency in order to be sent with Dr Wallich and Mr M'Clelland to visit and inspect the localities in which tea grew wild in Assam. Griffith's full report on this enquiry led to the important economic conclusion (based largely on a critical comparison of the Assam flora with the flora of tea-growing regions of China) that tea might be successfully grown under the conditions in Assam and similar districts of India. When the other members of the expedition returned Griffith was detained in Assam, where he remained during the whole of 1836, making a successful expedition into the Mishmee mountains only once before visited by a European.
Early in 1837 Griffith, accompanied by only one servant, set off on an exploring expedition through the very disturbed country of Burmah towards Rangoon. All news of him ceased, or rather his assassination was credited by the Government and reported in the newspapers, when in June he re-appeared, ragged and travel stained, in Calcutta. He had explored down the Hookhoom (Hokong) Valley and on to Ava, and had then proceeded more rapidly by river to Rangoon, conveying his collections with danger and difficulty.
Appointed Surgeon to the embassy about to start for Bhutan, he filled up the intervening two months by again going to the Khasi hills to collect. He then accompanied the expedition to Bhutan, traversing over four hundred miles of the country and returning to Calcutta in June 1838. Here he spent the next few months arranging his collections and also studying the plants of the suburbs.
In November he joined the army of the Indus and accompanied it in its whole march. He remained another year in Afghanistan making various expeditions in the country and into the Hindoo Koosh. He returned, after visiting Simla and the Nerbudda, to Calcutta in the middle of 1841.
Griffith then proceeded to Malacca where he had been appointed Civil Assistant-Surgeon. He remained only a year, but long enough to appreciate the great interest of the district for his botanical work and to complete some important observations. He collected the plants of the province and also visited Mount Ophir.
Recalled to Calcutta, he took charge of the Botanic Gardens and also lectured to the medical students during Wallich's absence from August 1842 to August 1844, pressing forward reforms in the gardens and using his opportunity for scientific observation. On Wallich's return Griffith remained for some months longer in Calcutta continuing his work, married in September, and returned to Malacca in December full of hopeful plans for scientific work there. He had barely arrived at Malacca and begun work than he was seized with a fatal illness and died on February 9, 1845.
It has been necessary to consider in some detail the rapid movements of Griffith's life in the East in order to fully appreciate the difficulties under which his large amount of scientific work was accomplished. The twelve years of his official life were filled with professional duties, difficult and dangerous exploration, management of the Botanic Gardens, and the labours entailed in making and caring for extensive collections. It would not have been surprising had Griffith, in spite of his attainments, contributed nothing to scientific botany beyond rendering these collections available for other workers. He estimated his collection of plants at more than twelve thousand species; and on his travels he did not neglect other collections of interest. Insects obtained by him are described, he collected the birds and fish in every district he visited; indeed he was a keen fisherman and must have thrown a fly in many a stream that had not been fished before, combining sport and science.
Griffith's collections were made with the definite purpose of enabling him, when he had leisure, to produce a general account of the Indian flora on a geographical basis. His methods of collecting were most enlightened and subserved his work as a morphologist and a student of the conditions of occurrence of the plants, not merely of formal systematic botany. The journals he kept on all expeditions are full of references to the occurrence of the plants met with. He often adopted a plan of roughly mapping each day's route and indicating the plants and associations of plants, along the line of march. I wonder if modern ecologists know of these records made long before ecology was invented?
Whenever possible he seems to have examined the morphology of the living plants, and he fully realised the value of preserving portions of the plants in spirit for future examination instead of relying on herbarium material.
This quotation from a letter to Wight (then Superintending Surgeon of the Madras Service), with whom Griffith kept up a most interesting and friendly correspondence, from which I should like to quote largely, may give an idea of his point of view and also show how he looked forward to returning to Malacca:—
"If ever you go to the place of Podostemon endeavour to get some germinating or at least very young plants. I can fancy how an Acotyledonous plant gets a stem but how a Dicotyledonous plant loses it, and becomes as some of them do, mere discs spread over rocks is another thing. Then again where are their roots? How opposed to late ideas of the absolute distinction of the three great divisions. Also please to take a bottle of spirits, and deposit specimens in it. I shall not be very sorry to get back to Malacca, this is a delightful place truly, but one is interrupted, and the lectures at the Medical College consume much time. For botany no place can exceed Malacca."
And again,
"What a business it will be to settle the types of the families from which the names must eventually be taken; this will never be done by dried-plant botanists; but by examination of development, which I am convinced will alone give the key."
As to Griffith's methods of work, we learn from a memorial notice of him by Mr M'Clelland that whenever possible after the business of the morning was finished the rest of the day was devoted "to the examination and dissection of plants under the microscope, drawing and describing all peculiarities presented." "Even on his death-bed his microscope stood beside him with the unfinished drawings and papers and dissections of plants on which he was engaged the day on which the fatal symptoms of his disorder came on."
All his work shows the same characters of direct individual observation and interpretation of the facts before him, repeated examination of the same point, and almost a prodigality of labour in recording his observations in drawings. At first under the influence of Robert Brown, he used the simple microscope with triplet lenses, but later he employed the compound microscope and in the year before his death writes hopefully of ordering a first-rate microscope when he obtains the arrears due to him from the Directors.
Griffith's high attainments were appreciated by the distinguished circle of English botanists of his time with whom he corresponded. Mr Solby, to whom he always sent home his papers for submission to the Linnean Society; Robert Brown, to whose work he constantly recurs with admiration, and whose judgment he trusted absolutely; Lindley; Sir William Hooker, who looked forward to his being settled permanently in charge of the Calcutta gardens, and Dr Wight may be named.
I may quote from a letter addressed to Griffith by von Martius of Munich, since it couples his own opinion and that of Robert Brown. "He (Brown) agrees with me in appreciating your spirited and enlightened investigations, and I now more than ever look forward to you as his successor—as the standard English botanist."
Only an outline of the nature of Griffith's scientific work with some details on selected subjects can be attempted here. His published works in the Transactions of the Linnean Society and elsewhere, important as they are, represent only a small fraction of his observations. But the wisdom and liberality of the East India Company has put us in possession of his unpublished notes and drawings (bequeathed with his collections to the Company) in the posthumously published volumes of Notulae ad plantas Asiaticas with the accompanying sets of plates. Though his papers were not ready or intended for publication in this form and suffer from having had to be arranged by another hand, they afford, together with his published work, a particularly good picture of how the problems of morphology and classification presented themselves to a keen investigator at this time.
Of his purely systematic work I shall not speak at length. In addition to smaller papers the most important contribution was his illustrated monograph on the Palms of British East India. In the Notulae numerous species are described and figured nearly always with reference to the morphology and physiology of the parts concerned. It is his investigations made with direct reference to morphology and reproduction that claim our attention most. In dealing with them it is convenient to treat of the main questions to which he directed his attention rather than of the separate papers. I shall call attention first to his work on the flower and on fertilisation in a number of plants, then to his observations on Cycas, and lastly to his work on the Cryptogams.
Interest in the structure of the ovule and the nature of fertilisation was widespread at the time Griffith worked. A few years previously Robert Brown had laid the foundations of the scientific study of the ovule and the behaviour of the pollen tube, and during Griffith's time the papers of Schleiden, which extended the comparative study of the ovule and advanced the important though erroneous view that the embryo originated inside the embryo-sac from the tip of the entering pollen tube, were appearing. Schleiden's text-book did not appear until too late to be known to Griffith. His interest was keen on continuing the work, that Brown had begun, on plants that only a resident in the tropics had the opportunity of studying properly, and the first volume of the Notulae, with the accompanying Icones, and the more systematic volume on the Monocotyledons and Dicotyledons contain his unpublished observations on the ovule and flowers of many plants.
His first paper in the Linnean Transactions was on the ovule of Santalum. Griffith observed and rightly interpreted the free prolongation of the embryo-sac from the nucellus, and described the application of the pollen tube to the summit of the embryo-sac, the development of the endosperm, and the origin and development of the embryo. He also recognised and figured the great prolongation backwards of the embryo-sac as an empty, absorbent caecum. At first he left the origin of the embryo doubtful, while recognising the advantages of the exposed embryo-sac for settling the question, but later he decided in favour of Schleiden's erroneous view that the embryo developed from the tip of the pollen tube. Griffith also examined the ovules of Osyris recognising the corresponding facts.
Comparison with the figures of Santalaceous ovules in Guignard's later work will serve to show both the magnificent accuracy in observation of Griffith and the limitation, running through all the work of the time, of not recognising the contents of the embryo-sac before fertilisation.
The Loranthaceae was another family on which the development of the embryo-sac and the processes of fertilisation and development of the fruit interested Griffith specially. Not only did he send his results home to the Linnean Society in two papers, but his descriptions and figures of all the species described in the Notulae take account of these morphological and developmental facts. He traced the development of the cavity of the ovary and regarded the ovules as reduced to their simplest expression—to an "amnios" or embryo-sac. And he observed the extension of the embryo-sacs up the style and the union of the pollen tube with the tip of the embryo-sac. His further description of the development of the embryo, endosperm and fruit is wonderfully exact if we allow for his regarding the long suspensor bearing the embryo as derived from the pollen tube growing down through the long embryo-sac.
Griffith thus recognised all the main peculiarities of Viscum and of Loranthus subsequently described more in detail in European species by Hofmeister (whose analysis of Griffith's work in 1859 is a great testimony to its accuracy) and later by Treub in the tropical species which had been studied by Griffith.
The Balanophoraceae was another group, on which Griffith made pioneer investigations. He collected and examined all the species he met with, partly from the systematic interest in supporting Robert Brown's objection to Lindley's class of Rhizantheae, but still more from his interest in the details of their reproduction. An examination of the plates from his memoirs, only published after his death, in the Linnean Transactions will show how fully he was aware of the structure of the archegonium-like female flower of Balanophora; of the relation of the pollen-grains and pollen tubes to it; and of the appearance of the endosperm which he mistook for the embryo. Throughout he compares the structure with the pistillum (archegonium) of Bryophyta.
Thus in the Balanophoraceae also Griffith laid the foundations on which the work of Hofmeister, and more recently that of Treub and Lotsy follow.
When at Malacca Griffith interested himself among many other problems in the ovule and the development of the seed of Avicennia. He had previously paid attention to the viviparous embryos of other Mangroves. This piece of work, when compared with Treub's re-examination of Avicennia, brings out so clearly Griffith's accuracy, so far as his means of observation allowed him to go, that we may look for a moment at how these two investigations, separated by forty years, compare.
Griffith recognised the development of the embryo-sac in the nucellus of the ovule which he took to be naked, missing the very slightly indicated integument. He followed the pollen tube to the tip of the embryo-sac and the development of the endosperm in its upper portion, where the embryo appeared. He saw the growth of the endosperm leading to its complete protrusion from the ovule and inverting the embryo so that its cotyledons point to the surface. Further he saw the long, empty, absorbent caecum grow out from the hinder end of the embryo-sac into the massive base of the young seed.
This account is substantially correct in all its facts, and Treub's work adds to it the cellular details of the origin of the embryo-sac, the setting apart of the endosperm cell to grow into the haustorium, and the details of segmentation of the embryo.
Such vivid, accurate, description of strange facts, when previous knowledge gave no clue, is in itself no mean scientific achievement.
To sum up Griffith's work on the morphology of the reproductive organs of the Angiosperms we see that he added many important facts and gave correct descriptions of what still remain among the most anomalous ovules and embryos. His methods did not enable him to distinguish clearly the contents of the embryo-sac, and he accepted and confirmed Schleiden's erroneous view of the origin of the embryo. But this hardly detracts from the directness and consequent value of all his observations.
Turning now to the Gymnosperms, we find again that Griffith devoted much attention to those forms that from his residence in the tropics he was in a position to study with most advantage. He describes in the Notulae his observations on the ovules and pollination of various Coniferae and Gnetaceae. But we may concentrate our interest on his work on Cycas. The rough structure of the young seed had already been described by Robert Brown who had recognised the gymnospermy of the group.
But Griffith's descriptions and figures are much more accurate—are indeed far in advance of those of much later observers—and add greatly to our knowledge of this plant. These two figures ([pl. xvi]) will speak for themselves and show how clearly Griffith had grasped the morphology of the Cycadean ovule, how faithfully he delineated the details, and how he sought in progressive development to throw light on the structure. He added to the previously imperfect description of the ovule an accurate account of the pollen chamber, and the proof that pollen grains entered and filled it. Further he followed the germination of the pollen grains, not merely recording the fact that the tubes penetrated the nucellus all around the pollen chamber, but ascertaining in how many days the tubes were put forth. His fullest description is unfortunately displaced in the Notulae under the heading of Thuja, but it is clear that it refers to the Cycas figured on the same plate as that plant.
From what has been said of the nature of Griffith's work on the ovules, both of Angiosperms and Gymnosperms, the complete omission of his name in recent works on the two groups that are in constant use is at least noteworthy.
Griffith was specially interested in the study of Cryptogamic plants. In a letter to Wight he says "I would like to be out with a work on Indian Cryptogamia of higher forms; so much so that if I see no chance of my succeeding to the Gardens, I intend sending away all my other collections, and devoting myself to this object and general development, which is obviously the keystone of the arch."
He left Algae and Fungi (with the exception of the Characeae) alone, and it is his work on the Bryophyta and Pteridophyta that concerns us. For information on his views on these plants we are dependent on his paper on Salvinia and Azolla and on the Notulae, put together as I have said from his notes after his death, and not intended for publication in this form. But there is no difficulty in getting a clear grasp of his point of view. This was a mistaken one—an attempt to bring into line the reproduction of the gametophyte of Bryophytes, the sporophyte of Vascular Cryptogams, and the flowering plant with its flower and fruit. It is easy to be wise after the event. In these comparisons Griffith belonged to his time with a much wider field of personal observation than most possessed.
Plate XVI
From Griffith's Notulae
Median section of the ovule of Cycas
Nucellar apex of Cycas with pollen chamber and pollen grains
We must bear in mind that at the time when Griffith worked no idea of the sexual and asexual alternating generations in Pteridophytes had been gained, although the prothallia had been observed preceding the growth of the plant in Equisetum and Ferns. It was not till some years after Griffith's death that fuller facts as to the sexual organs were obtained and led to the right comparisons.
Griffith's work on the Bryophyta shows the same power of observation as that on the ovule, but the difficulties due to imperfect instruments are more evident. His views on reproduction were here, however, clear, since the development of the capsule was definitely related to the fertilisation of the pistilla (archegonia) by the substance formed in the anthers. His figures indicate how much he saw, and how here also he sought in development the interpretation of mature structure.
His early interest in the Liverworts, especially the Marchantiaceae, continued, and all the forms he collected were carefully examined and figured with his usual accuracy.
One of the Liverworts Griffith described may be taken as an illustration to this part of our subject on account of the interest of its re-discovery and re-description in 1910 by Goebel. This is a plant collected in Assam and named Monosolenium tenerum. This Marchantiaceous plant is described as having no air-chamber layer, as bearing sessile, dorsal, antheridial receptacles, and terminal, shortly stalked archegoniophores with one ventral groove in the stalk. A single archegonium—later capsule—is found in each of the half-dozen involucres. Spores and irregular bodies were found in the capsule.
Recently Goebel had two tea-plants sent home from Canton. They died, but he kept the soil moist on the chance of germinating seeds. Among a number of other plants there turned up a new Liverwort. On examination this proved to be Griffith's Monosolenium—all types of which had been lost—a most interesting form related to the Corsiniaceae.
In the Mosses and the Liverworts generally Griffith was clear on the development of the capsule or fruit following on the impregnation of an archegonium. But in Anthoceros, while he recognised the antheridia he was not clear as to the sunken archegonia, and regarded the capsules as arising by impregnation of unrecognisable spots on the young frond or thallus. He observed however the indication of the canal of the archegonial neck above the young capsule.
Analogy with Anthoceros confirmed him in his views on the reproduction of ferns. Here he spent much labour in considering the view, originally due to Hedwig, that the ramenta were male organs by the effect of which the sporangia developed. Griffith saw that if this was so, since the sporangia are initiated very early, the only time to search for the male organs was in the very young stage of the leaf. On examining such young leaves he found the terminal cells of the young ramenta very prominent and formed the working hypothesis that they were the male organs. But he stated this cautiously and was well aware how imperfect his means of observation were.
The whole line of work brings vividly before us how cryptogamic the Cryptogams were at this period.
Without attempting to survey Griffith's views on the various groups of Vascular Cryptogams, a word must be said of those on Salvinia and Azolla, on which he published a long paper in addition to the other descriptions and figures in the Notulae. His observations bear on the development of the sorus and sporangium, but he dismissed the microsporangia as abortive or imperfectly developed structures. (I may note in passing that the study of their development led him to regard the microsporangia of Isoetes in the same way.) He dwelt on the similarity of the sporangium and indusium of Azolla to a gymnospermus ovule, and regarded the filaments of Anabena seen penetrating within the indusium as probably the fertilising bodies in this naked-seeded cryptogam.
Thus with a large amount of fresh and original observation Griffith was on wrong lines in his general views and comparison—he classed the higher Cryptogams in his Notulae as
| Pistilligerous. | Musci. | Hepaticae. | ||
| Gymnospermous. | Azolla. | Salvinia. | Chara. | |
| Cryptogamous. | Ferns. | Lycopods. | Isoetes. | Marsilidae. |
| Anthocerotidae. Equisetidae. | ||||
Griffith's general views of the reproduction of all the Vascular Cryptogams was necessarily wrong, since the prime clue of the recognition of the prothallus and plant as distinct had not been found. In this connection his figuring young plants of Equisetum attached to prothalli is interesting. In some speculations concerning the embryology of Loranthus he came, by a wrong line of approach, within touch of the right comparison, when he compares the endosperm to the confervoid green growth (i.e. the prothallus) at the base of the young plant of Equisetum.
It is idle to speculate on what might have happened had such a wide observer as Griffith chanced on the clue. In this respect he was of his time as most are. The man who put the industrious but blind gropings of this period in morphological botany straight, both as regards the development of the embryo and the comparative ontogeny of archegoniate plants was Hofmeister, and like all exceptional men he belonged to the new period created by him.
The great advantage of this course of lectures seems to me to be that it approaches the study of the history of botany in the right way; for progress in our science has been the result of individuals rather than of schools. The consideration of the work of Griffith from 1832 to 1845 is a vivid illustration of the condition of morphological botany in the earlier portion of the period, surveyed in one of the chapters in Sach's History under the title of "Morphology and Systematic Botany under the influence of the History of Development and the knowledge of the Cryptogams." These two subjects were always before Griffith.
The interest of the personality of William Griffith and of the work he accomplished in his tragically short life is obvious. Not less so is the way in which that work was done inside the limitations of his period. We, who are still gleaners in the field that Griffith and his contemporaries cleared and Hofmeister marked out and tilled, are probably just as incapable of conceiving the future developments of morphology.
[ARTHUR HENFREY]
1819-1859
By F. W. OLIVER
Narrative—state of Botany—dawn of the Golden Age—sexuality of Angiosperms—Schleiden's elucidation of fern life-history—Nägeli, Suminski and Hofmeister—recognition by Henfrey—original work—publications—the Micrographic Dictionary—The Botanical Gazette—its features—Henfrey's labours not immediately productive.
The claim of Henfrey to rank among the founders of botany in this country depends less on his own original contributions than on a whole-hearted devotion to the propagation and diffusion of the newer methods and results which marked an epoch during the forties and fifties of last century. The outset of Henfrey's career coincided with a great turning point in the history of botany, and to Henfrey will always belong the credit of being the first Englishman to recognise the full significance of the movement. From that moment he unceasingly made known and diffused in this country the results of the German renaissance. That Henfrey should have failed to establish the newer botany in England was the result of a variety of circumstances, one of which was his early death.
The available biographical material of Henfrey being extremely meagre, it has been necessary in preparing the present account to rely almost entirely on his published writings. In some ways this lack of personal details is no disadvantage as our present interest in Henfrey depends essentially on the movement in botany with which he was identified.
Arthur Henfrey was born at Aberdeen, in 1819, of English parents. He underwent the usual course of training for the medical profession at St Bartholomew's Hospital—becoming a member of the Royal College of Surgeons in 1843. In consequence of bronchial trouble, to which he eventually succumbed at the early age of 39, Henfrey never practised his profession. Compelled to a life of seclusion he at once turned to a scientific career and more particularly to the pursuit of botany. In 1847 he undertook the duties of Lecturer in Botany at St George's Hospital Medical School, where among his colleagues was Edwin Lankester, himself a redoubtable naturalist and the father of Sir Ray Lankester, the eminent zoologist of our own day.
Henfrey succeeded Edward Forbes as Professor of Botany in King's College, London, in 1852—a post which he held till his death. He was elected to the Fellowship of the Royal Society in the same year.
He died quite suddenly in 1859, at the house on Turnham Green, where he had resided for many years.
In order to understand the part played by Henfrey, it is necessary briefly to review the state of botany in the first half of the nineteenth century.
Linnaeus of course, botanically, the outstanding fact of the eighteenth century, was no exception to the dictum that "the evil that men do lives after them."
It was supposed that botany had reached its culminating point in Linnaeus and that improvement could only be made in details—elaborating and extending his system. As Sachs tells us in his History, the result was that "Botany ceased to be a science; even the describing of plants which Linnaeus had raised to an art became once more loose and negligent in the hands of his successors. Botany gradually degenerated under the influence of his authority into an insipid dilettantism—a dull occupation for plant collectors who called themselves systematists, in entire contravention of the meaning of the word."
This was written with especial reference to Germany, but it applied with no less force to our own country where the Linnaean idea had taken deep root and the Linnaean collections had found a sanctuary.
However, by 1840, a change was coming over the face of botany. Little as it can have been dreamt, the Golden Age was already beginning—destined in a relatively short time to transform the subject. This Golden Age was contemporaneous with, and immediately dependent on, the rise of a group of young botanists in the Fatherland, a group which included von Mohl, Schleiden, Hofmeister, Nägeli, Cohn and De Bary. Later it was reinforced by Sachs, who in addition to being a brilliant physiologist was a gifted writer who did much to establish scientific botany on a sound footing. It is impossible to overestimate the debt due to Sachs, particularly for his great Textbook of Botany, which at the right psychological moment brought the whole of the modern work between the covers of a single volume.
It was with the dawn of this period that Henfrey identified himself. In the 15 years of his active career (1844-1859) he devoted himself very largely to making his fellow-countrymen acquainted with the newer aspects of botany. More particularly it was the recent discoveries as to the reproduction and life history of the Vascular Cryptogams that specially engaged his interest—the researches which broadly speaking we associate with Hofmeister to-day.
Before we go on to speak of the sexuality of the Cryptogams however, a few words may be devoted to that of the flowering plants.
Sexuality of Flowering Plants. At the period when Henfrey entered on his career as a botanist no reasonable doubt remained as to the existence of sexes among the flowering plants. The theory of the sexual significance of the organs of the flower, brilliantly founded by Koelreuter in the previous century, had been perfected with a great volume of experimental proof by K. F. Gaertner the son of Joseph Gaertner of Carpologia fame.
By 1830 the mechanism of fertilisation came to light in Amici's discovery of the pollen tube which he traced from the stigma to the micropyle. The microscopic aspect of the problem was taken up with great energy by Schleiden and brought to the forefront as the burning question of the early forties. The theory of Schleiden, which applied in particular to the flowering plants, made its influence felt to such an extent in the search for evidence of sexuality among the Cryptogams, that we may conveniently state in a few words in what it consisted.
Schleiden traced the pollen tube into the micropyle, and thence to the nucellus where it depressed or invaginated the apex of the embryo-sac, and in the recess or indentation so produced the tip of the pollen tube was converted into the embryo—its actual apex being represented by the plumule. This theory was the lineal descendant in modernised trappings of the old view expressed by Morland and others at the beginning of the eighteenth century that the embryo was contained in the pollen grain, and that the ovule was no more than the brood chamber whither it must be brought to undergo further development. This erroneous interpretation of the true facts was always repudiated by Amici, and was finally overthrown by Hofmeister and Radlkofer in the early fifties. In this connection we may note in passing Henfrey's careful paper on the impregnation of Orchis Morio, published in 1856, which fully corroborated Amici. In this paper the relations of pollen tube, embryo-sac, egg-cell, suspensor and embryo were correctly interpreted, and the new point established, contrary to the assertions of previous observers, that the ovum or "germinal-vesicle," prior to fertilisation, was a naked, unwalled cell.
Sexuality in Cryptogams. By far the most important question that came to a head in Henfrey's time was that of the morphological relationships of the Cryptogams and flowering plants. Hitherto these had remained altogether obscure in the absence of reliable data based on the proper application of the microscope to the elucidation of the life histories of the lower plants. Under the influence of the Linnaean school, which had taken deep root in this country, as elsewhere, the systematic study of flowering plants had been widely pursued, and in so far as the ferns were concerned their homologies were commonly interpreted in terms of the flowering plants. Without any real guidance in fact, a great diversity of views of these homologies found expression. The following, taken from Lindley, may serve to illustrate their general nature.
The sorus was regarded as a sort of compound fruit, the sporangium as a carpel, the annulus as its midrib, and the spores as the seeds. Speculations such as these are of the same order as the crude conjectures which with less excuse relieve the answer books of examination candidates at the present time.
In the search for the male organs of the fern attention was naturally directed to the neighbourhood of the sorus, and the stomata, indusia and glandular appendages were in turn mistaken by various observers for the anthers. The "limit" was reached by Griffith who, as is stated at [page 190], conjectured that the Anabena filaments which accompany the megasporangia of Azolla were no other than the male organs of that plant.
Schleiden spoke of these researches with the utmost scorn. "For my part I am surprised that no one has yet insisted upon the presence of the organs of sense, as eyes and ears in plants, since they are possessed by animals. Such an assumption would not be a bit more absurd than the mania of insisting upon having anthers in the Cryptogams, simply because they are found in the Phanerogams."
All these ill-grounded hypotheses were swept away in 1844 when Nägeli discovered antheridia containing spermatozoids on the "cotyledon" or pro-embryo of the fern—the prothallus we call it now. Nägeli at once recognised their essential agreement with the antheridia already known in the Bryophytes and compared the spermatozoids with the corresponding structures in animals. But as he overlooked the existence of the archegonia, or rather by some lapse mistook them for stages in the development of the antheridia, it is not surprising that he was at a loss to understand the significance of his discovery, and that he should have commented on his dilemma in the following terms. "Seeing that the female organs (spores) arise on the frond at a much later stage of development, and long after the pro-embryo has died away, the function of the spermatozoids is far from evident."
It was only three years later that light was thrown on the situation, and from an unexpected quarter. Count Suminski, an amateur microscopist, announced the discovery of additional reproductive organs on the fern pro-embryo, which he clearly distinguished from the "spiral filament organs" or antheridia. His full paper, which appeared in 1848, marks an epoch in morphology, and was a very remarkable performance. In it he redescribes the antheridia and spermatozoids—detecting their tufted cilia which Nägeli had overlooked. The archegonia he describes as ovules without envelopes consisting of a papilla (the neck) which becomes perforated, giving the spermatozoid access to the embryo-sac within. His figures of the process of fertilisation are extremely interesting as they show how completely he was dominated by the theory of Schleiden to which allusion has already been made. The head of the sperm is represented as entering the "embryo-sac," and there becoming encysted to form the embryo just as the tip of the pollen tube was supposed to do in flowering plants. The further development of the embryo and its various organs are traced and figured, however, in the most admirable way. At the conclusion of his paper Suminski states that in view of the presence of male organs and ovules, and the occurrence of fertilisation, the cryptogamy of ferns does not exist in a physiological sense, and ceases to have any validity as a peculiar character. A remark which he follows up by the statement that ferns must on the existing classification be referred to the Monocotyledons.
In certain respects no doubt Suminski's paper is fantastic—more especially the circumstantial details given of the process of fertilisation. But, however we may criticise his work the credit belongs to Suminski of showing (1) that sexual organs are borne on the prothallus, (2) that the embryo fern plant is produced as the result of fertilisation. Unlike Nägeli, to Suminski came the happy inspiration of looking for the female organs in the position where common sense indicated they ought to be found.
Suminski's paper instantly aroused universal interest, and the whole of his assertions were at first categorically denied by the German botanist Wigand.
We may now trace Henfrey's attitude to Suminski's work.
His first notice occurs in the body of a review of Lindley's "Introduction" in the first volume of his Botanical Gazette, and shows him to have been profoundly sceptical, if not contemptuous, of the occurrence of fertilisation in the prothallus of the fern. His words are "this (i.e. Suminski's discovery) appears to have little but originality to render it worthy of notice." That appeared in February 1849.
Writing at greater length of Suminski's work in the Annals and Magazine of Natural History, in November of the same year, he speaks much more guardedly. "These researches are in the highest degree curious, and if the facts related prove to be correct, most importantly affect the received views of analogies in the generative processes of plants."
At the same time Henfrey says he hopes to speak more definitely on this matter when his own investigations are complete. Two years later his own very careful work in the same field was laid before the Linnean Society, in which he corroborated the main facts that had come to light. Turning once again to the paper of Suminski, after making certain criticisms of detail, Henfrey handsomely remarks—"Nothing however can take from him the credit of having discovered the archegonia and their import, one of the most important discoveries in physiological Botany of modern times since it has led to results revolutionising the whole theory of the reproduction of plants and opened out a totally new sphere of inquiry into the laws and relations of vegetable life."
For some little time after these discoveries the archegonia of the fern were, on the initiative of Mercklin, commonly referred to as the "organs of Suminski," a custom which happily fell into desuetude. Mercklin, in his paper, which essentially repeats the work of Suminski, states that he devoted his entire attention for three months to the fern prothalli before he succeeded in observing the entrance of a spermatozoid.
In reviewing the early papers of the Hofmeisterian epoch—papers which form the bed-rock of the existing morphology—one is struck with the marvellous rapidity with which their significance was apprehended. We find the phrase "alternation of generations" employed within two years of the discoveries of Suminski, whilst by the early fifties the general genetic relations of the vascular series were realized in quite a new light.
As Sachs puts it:—"When Darwin's theory was given to the world eight years after Hofmeister's investigations, the relations of affinity between the great divisions of the vegetable kingdom were so well established and so patent that the theory of descent had only to accept what genetic morphology had actually brought to view."
Among Henfrey's original contributions other than those dealing with the burning questions already mentioned, was a series dealing with the Anatomy of Monocotyledons. This would appear to have led him on to study the Nymphaeaceae, and especially the anatomy of Victoria regia—a paper which may be compared perhaps with Prof. Gwynne-Vaughan's more recent study. Henfrey was quite alive to the monocotyledonous affinity, and the enlightened and, for that date, unconventional views to which he gave expression, drew an interesting notice by Hooker and Thomson in the first volume of their Indian Flora.
Another of his papers dealt rather fully with the development of the spores and elaters of Marchantia, where he filled in a considerable lacuna in the knowledge of that group. It is curious to find as late as 1855 so intelligent and well informed a botanist as Henfrey laying it down that the cells of Marchantia, in particular, and Liverworts in general, were destitute of nuclei. It is superfluous to say that this apprehension was quite baseless. Indeed, forty years later, the group of the Liverworts was deliberately chosen by Prof. J. B. Farmer, for the investigation of nuclear phenomena on account of the favourable conditions under which they could be studied!
Microtechnique at that time was of course a much simpler affair than it has since become. Contemporary papers as a rule say little about methods; however one of Henfrey's occasional notes in a magazine tells us that caustic potash, iodine, sulphuric, hydrochloric and acetic acids, together with ether were in common use. Schultze's reagent—chloride of zinc iodide—was invented in 1850, but does not appear to have been generally employed till many years later.
It would however be a serious error to underestimate the value of the earlier work in plant histology. The present writer once spent an interesting morning in Pfeffer's laboratory at Tübingen rummaging through hundreds of the great von Mohl's anatomical preparations. Among these were sections of palm endosperms in which the, at that time recently discovered, continuity of the protoplasm through the cell walls was plainly visible. The existence of these filaments had been detected by von Mohl some years before, but he had refrained from publishing his observations from over-cautiousness.
As a translator and editor Henfrey was responsible for the English edition of von Mohl's Principles of the Anatomy and Physiology of the Vegetable Cell, published in 1852, for two volumes of Reports on Botany in the Ray Society's publications, whilst he had a considerable share in Lankester's translation of Schleiden's famous Principles of Scientific Botany, 1847. In addition to these there were constant abstracts and critical reviews from his pen in the Annals and Magazine of Natural History—a journal of which he became botanical editor before the close of his life.
As a writer of text-books Henfrey was very prolific. First came his Outlines of Botany, 1847, followed by the Rudiments of Botany. Much more ambitious was his Elementary Course of Botany which became a standard text-book running through numerous editions after his death, under the editorship of the late Dr M. T. Masters. To these must be added, in conjunction with Griffith[106], the Micrographic Dictionary, a substantial volume dealing in innumerable special and general articles with the microscopic study of plants and animals. This work was no mere compilation, but embodied in its pages is a very large amount of independent observation. The illustrations covering nearly fifty plates were by Tuffen West, and reached a high degree of excellence. A well known botanist, a contributor to the present volume, has more than once assured me that it was to the Micrographic Dictionary that he owed his salvation!
Should anyone desire to get a vivid and accurate picture of the precise state of Botany in this country at the middle of the last century, he cannot do better than turn over the pages of The Botanical Gazette, a monthly journal of the progress of British botany, founded and conducted by Henfrey. It was about the size of our own New Phytologist, with which it had not a little in common. In one respect it differed; unlike the New Phytologist the Gazette was financially a failure and after carrying it on at his own expense for three years (1849-1851) Henfrey had to relinquish the undertaking.
A perusal of its contents clearly shows that its editor regarded his journal as one of the instruments of diffusing the New Botany. Having to rely largely for his subscribers upon the amateur collector he points out in the prefatory note that a feature will be made not only of home botany but also of contributions or abstracts from abroad dealing with floras which have much in common with our own. For the benefit of those whose collections had reached considerable dimensions, and for whom the lack of new plants might connote a waning stimulus, he held out the further inducement of papers on the general anatomy of familiar plants, of which an excellent example by Thilo Irmisch on the stolons of Epilobium was included in the first number.
For the three years of its existence Henfrey kept faith with the British botanists and a number of The Botanical Gazette rarely appeared without an article contrived for their edification. The task was evidently a congenial one, for Henfrey had a sound knowledge of British plants with especial reference to geographical distribution and critical forms. Unlike several later exponents of the New Botany, Henfrey was quite able to hold his own with the systematists. He more than once expresses the opinion that there was too great a tendency to lump species in the handbooks to the Flora, and he urged on the occasion of the preparation of the third edition of the London Catalogue of British Plants that many more forms should find recognition. The editors of the catalogue however successfully opposed the suggestion on the ingenious grounds that it would raise the weight for postage beyond the limits of a blue (twopenny) stamp!
Henfrey thought much might be done by cultivation under varying conditions to settle vexed questions as to critical species, and suggested that a limited number of botanists in different parts of the country should co-operate in a scheme under which seed should annually be distributed, harvested and re-distributed among those taking part. Henfrey himself offered to undertake the somewhat onerous duty of receiving and distributing the seed and of generally correlating the work. As however his proposal was merely tagged on to a note on Sagina apetala and ciliata it is hardly remarkable that nothing came of it.
An interesting minor feature of the Gazette was the reporting of the proceedings of the various Botanical Societies throughout the country. These show that a chronic state of intellectual famine frequently obtained even at the leading societies—a state of which vestiges are still occasionally discernible. It was no unusual occurrence at the Linnean even during the period of Robert Brown's presidency for the meeting to be regaled with long extracts from the commentaries on the Hortus Malabaricus. In this respect however the record was easily held by the now defunct Botanical Society of London, which eked out its programme for a whole year with a communication by a Mr D. Stock "On the Botany of Bungay, Suffolk." Begun on the 11th October, 1850, it only drew to a conclusion on the 3rd October, 1851. There were other attractive features in The Botanical Gazette on which space does not allow me to dwell.
The general impression gained, however, from a perusal of the papers of that time is that they were refreshingly short, as compared with our own day, and often very much to the point. The recording of observations was rarely made the occasion for a survey of the whole field of botany, and little trace was discernible of the present habit of over-elaboration.
The foregoing outline of Henfrey's activities shows that they were devoted wholly to the spread of the Newer Botany in this country. The means employed included the publication of reviews and abstracts, the editing of translations of the more notable books, the founding of journals, and the writing of text-books. Moreover by his own investigations he kept close touch with the modern work and was indeed the means of corroborating and often materially advancing many of the larger problems before putting them into general circulation in this country.
And yet, in spite of this complete devotion of his life to the cause, the New Botany found no permanent place in this country till twenty years after Henfrey's death.
Botanically speaking, the organisation and rise of taxonomy was the ruling pre-occupation of the period under consideration, a direct outcome of colonial expansion and consolidation. Fed on unlimited supplies of new material from the ends of the earth the taxonomic habit became supreme. What could an isolated student and recluse like Henfrey do to stem this flood? Circumstances were too strong for him, and founding no immediate school it remained for a later generation to take up the task.
Though the history of the establishment of the New Botany in England lies outside the province of this lecture, it is instructive, as a contrast in methods, to note the manner of its accomplishment. Henfrey, who relied on his pen, had proved ineffective to bring about a revolution. Twenty years later it fell to Sir William Thiselton-Dyer, then a young man, to succeed where Henfrey had failed. By his enlightened teaching and personal magnetism, Thiselton-Dyer aroused a widespread interest in laboratory botany. But the matter was not allowed to rest there. Holding as he did an important post at Kew, the strategic centre, he was able to obtain appointments in the chief Colleges and Universities of the country for the recruits whom he had attracted. In this way, by the exercise of an acute intelligence amounting to statesmanship, and in a very short period of time, the New Botany became everywhere firmly established.