Cycas revoluta ([see page 311]).

(Frontispiece.)

ELEMENTARY BOTANY

BY

GEORGE FRANCIS ATKINSON, Ph.B.

Professor of Botany in Cornell University

THIRD EDITION, REVISED

NEW YORK
HENRY HOLT AND COMPANY
1905

Copyright, 1898, 1905
BY
HENRY HOLT AND COMPANY

ROBERT DRUMMOND, PRINTER, NEW YORK

PREFACE.

The present book is the result of a revision and elaboration of the author’s “Elementary Botany,” New York, 1898. The general plan of the parts on physiology and general morphology remains unchanged. A number of the chapters in the physiological part are practically untouched, while others are thoroughly revised and considerable new matter is added, especially on the subjects of nutrition and digestion. The principal chapters on general morphology are unchanged or only slightly modified, the greatest change being in a revision of the subject of the morphology of fertilization in the gymnosperms and angiosperms in order to bring this subject abreast of the discoveries of the past few years. One of the greatest modifications has been in the addition of chapters on the classification of the algæ and fungi with studies of additional examples for the benefit of those schools where the time allowed for the first year’s course makes desirable the examination of a broader range of representative plants. The classification is also carried out with greater definiteness, so that the regular sequence of classes, orders, and families is given at the close of each of the subkingdoms. Thus all the classes, all the orders (except a few in the algæ), and many of the families, are given for the algæ, fungi, mosses, liverworts, pteridophytes, gymnosperms, and angiosperms.

But by far the greatest improvement has been in the complete reorganization, rewriting, and elaboration of the part dealing with ecology, which has been made possible by studies of the past few years, so that the subject can be presented in a more logical and coherent form. As a result the subject-matter of the book falls naturally into three parts, which may be passed in review as follows:

Part I. Physiology. This deals with the life processes of plants, as absorption, transpiration, conduction, photosynthesis, nutrition, assimilation, digestion, respiration, growth, and irritability. Since protoplasm is fundamental to all the life work of the plant, this subject is dealt with first, and the student is led through the study of, and experimentation with, the simpler as well as some of the higher plants, to a general understanding of protoplasm and the special way in which it enables the plant to carry on its work and to adjust itself to the conditions of its existence. This study also serves the purpose of familiarizing the pupil with some of the lower and unfamiliar plants.

Some teachers will prefer to begin the study with general morphology and classification, thus studying first the representatives of the great groups of plants, and others will prefer to dwell first on the ecological aspects of vegetation. This can be done in the use of this book by beginning with Part II or with Part III.

But the author believes that morphology can best be comprehended after a general study of life processes and functions of the different parts of plants, including in this study some of the lower forms of plant life where some of these processes can more readily be observed. The pupil is then prepared for a more intelligent consideration of general and comparative morphology and relationships. Even more important is a first study of physiology before taking up the subject of ecology. The great value to be derived from a study of plants in their relation to environment lies in the ability to interpret the different states, conditions, behavior, and associations of the plant, and for this physiology is indispensable. It is true that a considerable measure of success can be obtained by a good teacher in beginning with either subject, but the writer believes that measure of success would be greater if the subjects were taken up in the order presented here.

Part II. Morphology and life history of representative plants. This includes a rather careful study of representative examples among the algæ, fungi, liverworts, mosses, ferns and their allies, gymnosperms and angiosperms, with especial emphasis on the form of plant parts, and a comparison of them in the different groups, with a comparative study of development, reproduction, and fertilization, rounding out the work with a study of life histories and noting progression and retrogression of certain organs and phases in proceeding from the lower to the higher plants. Thus, in the algæ a first critical study is made of four examples which illustrate in a marked way progressive stages of the plant body, sexual organs, and reproduction. Additional examples are then studied for the purpose of acquiring a knowledge of variations from these types and to give a broader basis for the brief consideration of general relationships and classification.

A similar plan is followed in the other great groups. The processes of fertilization and reproduction can be most easily observed in the lower plants like the algæ and fungi, and this is an additional argument in favor of giving emphasis to these forms of plant life as well as the advantage of proceeding logically from simpler to more complex forms. Having also learned some of these plants in our study of physiology, we are following another recognized rule of pedagogy, i.e., proceeding from known objects to unknown structures and processes. Through the study of the organs of reproduction of the lower plants and by general comparative morphology we have come to an understanding of the morphology of the parts of the flower, and of the true sexual organs of the seed plants, and no student can hope to properly interpret the significance of the flower, or the sexual organs of the seed plants who neglects a careful study of the general morphology of the lower plants.

Part III. Plant members in relation to environment. This part deals with the organization of the plant body as a whole in its relation to environment, the organization of plant tissues with a discussion of the principal tissues and a descriptive synopsis of the same. This is followed by a complete study from a biological standpoint of the different members of the plant, their special function and their special relations to environment. The stem, root, leaf, flower, etc., are carefully examined and their ecological relations pointed out. This together with the study of physiology and representatives in the groups of plants forms a thorough basis for pure plant ecology, or the special study of vegetation in its relation to environment.

There is a study of the factors of environment or ecological factors, which in general are grouped under the physical, climatic, and biotic factors. This is followed by an analysis of vegetation forms and structures, plant formations and societies. Then in order are treated briefly forest societies, prairie societies, desert societies, arctic and alpine societies, aquatic societies, and the special societies of sandy, rocky, and marshy places.

Acknowledgments. The author wishes to express his gratefulness to all those who have given aid in the preparation of this work, or of the earlier editions of Elementary Botany; to his associates, Dr. E. J. Durand, Dr. K. M. Wiegand, and Professor W. W. Rowlee, of the botanical department, and to Professor B. M. Duggar of the University of Missouri, Professor J. C. Arthur of Purdue University, and Professor W. F. Ganong of Smith College, for reading one or more portions of the text; as well as to all those who have contributed illustrations.

Illustrations. The large majority of the illustrations are new (or are the same as those used in earlier editions of the author’s Elementary Botany) and were made with special reference to the method of treatment followed in the text. Many of the photographs were made by the author. Others were contributed by Professor Rowlee of Cornell University; Mr. John Gifford of New Jersey; Professor B. M. Duggar, University of Missouri; Professor C. E. Bessey, University of Nebraska; Dr. M. B. Howe, New York Botanical Garden; Mr. Gifford Pinchot, Chief of the Bureau of Forestry; Mr. B. T. Galloway, Chief of the Bureau of Plant Industry; Professor Tuomey of Yale University; and Mr. E. H. Harriman, who through Dr. C. H. Merriam of the National Museum allowed the use of several of his copyrighted photographs from Alaska. To those who have contributed drawings the author is indebted as follows: to Professor Margaret C. Ferguson, Wellesley College; Professor Bertha Stoneman of Huguenot College, South Africa; Mr. H. Hasselbring of Chicago; Dr. K. Miyake, formerly of Cornell University and now of Doshisha College, Japan; and Professors Ikeno and Hirase of the Tokio Imperial University. The author is also indebted to Ginn & Co., Boston, for the privilege to use from his “First Studies of Plant Life” the following figures: 28, 29, 46, 48, 49, 56, 62, 66, 67, 87, 102, 103, 422-426, 429, 430, 438-440, 443, 444, 448, 449, 452, 472-475. A few others are acknowledged in the text.

Cornell University, April, 1905.

TABLE OF CONTENTS.

PART I.
PHYSIOLOGY.

[CHAPTER I.]
PROTOPLASM. [1]

1. In the study of plant life and growth, it will be found convenient first to inquire into the nature of the substance which we call the living material of plants. For plant growth, as well as some of the other processes of plant life, are at bottom dependent on this living matter. This living matter is called in general protoplasm.

2. In most cases protoplasm cannot be seen without the help of a microscope, and it will be necessary for us here to employ one if we wish to see protoplasm, and to satisfy ourselves by examination that the substance we are dealing with is protoplasm.

3. We shall find it convenient first to examine protoplasm in some of the simpler plants; plants which from their minute size and simple structure are so transparent that when examined with the microscope the interior can be seen.

For our first study let us take a plant known as spirogyra, though there are a number of others which would serve the purpose quite as well, and may quite as easily be obtained for study.

Protoplasm in spirogyra.

4. The plant spirogyra.—This plant is found in the water of pools, ditches, ponds, or in streams of slow-running water. It is green in color, and occurs in loose mats, usually floating near the surface. The name “pond-scum” is sometimes given to this plant, along with others which are more or less closely related. It is an alga, and belongs to a group of plants known as algæ. If we lift a portion of it from the water, we see that the mat is made up of a great tangle of green silky threads. Each one of these threads is a plant, so that the number contained in one of these floating mats is very great.

Let us place a bit of this thread tangle on a glass slip, and examine with the microscope and we will see certain things about the plant which are peculiar to it, and which enable us to distinguish it from other minute green water plants. We shall also wish to learn what these peculiar parts of the plant are, in order to demonstrate the protoplasm in the plant.[2]

5. Chlorophyll bands in spirogyra.—We first observe the presence of bands; green in color, the edges of which are usually very irregularly notched. These bands course along in a spiral manner near the surface of the thread. There may be one or several of these spirals, according to the species which we happen to select for study. This green coloring matter of the band is chlorophyll, and this substance, which also occurs in the higher green plants, will be considered in a later chapter. At quite regular intervals in the chlorophyll band are small starch grains, grouped in a rounded mass enclosing a minute body, the pyrenoid, which is peculiar to many algæ.

Fig. 1.

Thread of spirogyra, showing long cells, chlorophyll band, nucleus, strands of protoplasm, and the granular wall layer of protoplasm.

6. The spirogyra thread consists of cylindrical cells end to end.—Another thing which attracts our attention, as we examine a thread of spirogyra under the microscope, is that the thread is made up of cylindrical segments or compartments placed end to end. We can see a distinct separating line between the ends. Each one of these segments or compartments of the thread is a cell, and the boundary wall is in the form of a cylinder with closed ends.

7. Protoplasm.—Having distinguished these parts of the plant we can look for the protoplasm. It occurs within the cells. It is colorless (i.e., hyaline) and consequently requires close observation. Near the center of the cell can be seen a rather dense granular body of an elliptical or irregular form, with its long diameter transverse to the axis of the cell in some species; or triangular, or quadrate in others. This is the nucleus. Around the nucleus is a granular layer from which delicate threads of a shiny granular substance radiate in a starlike manner, and terminate in the chlorophyll band at one of the pyrenoids. A granular layer of the same substance lines the inside of the cell wall, and can be seen through the microscope if it is properly focussed. This granular substance in the cell is protoplasm.

8. Cell-sap in spirogyra.—The greater part of the interior space of the cell, that between the radiating strands of protoplasm, is occupied by a watery fluid, the “cell-sap.”

9. Reaction of protoplasm to certain reagents.—We can employ certain tests to demonstrate that this granular substance which we have seen is protoplasm, for it has been found, by repeated experiments with a great many kinds of plants, that protoplasm gives a definite reaction in response to treatment with certain substances called reagents. Let us mount a few threads of the spirogyra in a drop of a solution of iodine, and observe the results with the aid of the microscope. The iodine gives a yellowish-brown color to the protoplasm, and it can be more distinctly seen. The nucleus is also much more prominent since it colors deeply, and we can perceive within the nucleus one small rounded body, sometimes more, the nucleolus. The iodine here kills and stains the protoplasm. The protoplasm, however, in a living condition will resist for a time some other reagents, as we shall see if we attempt to stain it with a one per cent aqueous solution of a dye known as eosin. Let us mount a few living threads in such a solution of eosin, and after a time wash off the stain. The protoplasm remains uncolored. Now let us place these threads for a short time, two or three minutes, in strong alcohol, which kills the protoplasm. Then mount them in the eosin solution. The protoplasm now takes the eosin stain. After the protoplasm has been killed we note that the nucleus is no longer elliptical or angular in outline, but is rounded. The strands of protoplasm are no longer in tension as they were when alive.

Fig. 2.
Cell of spirogyra before
treatment with iodine.

Fig. 3.
Cell of spirogyra after
treatment with alcohol and iodine.

10. Let us now take some fresh living threads and mount them in water. Place a small drop of dilute glycerine on the slip at one side of the cover glass, and with a bit of filter paper at the other side draw out the water. The glycerine will flow under the cover glass and come in contact with the spirogyra threads. Glycerine absorbs water promptly. Being in contact with the threads it draws water out of the cell cavity, thus causing the layer of protoplasm which lines the inside of the cell wall to collapse, and separate from the wall, drawing the chlorophyll band inward toward the center also. The wall layer of protoplasm can now be more distinctly seen and its granular character observed.

We have thus employed three tests to demonstrate that this substance with which we are dealing shows the reactions which we know by experience to be given by protoplasm. We therefore conclude that this colorless and partly granular, slimy substance in the spirogyra cell is protoplasm, and that when we have performed these experiments, and noted carefully the results, we have seen protoplasm.

Fig. 4.

Cell of spirogyra before
treatment with glycerine.

Fig. 5.

Cells of spirogyra after
treatment with glycerine.

11. Earlier use of the term protoplasm.—Early students of the living matter in the cell considered it to be alike in substance, but differing in density; so the term protoplasm was applied to all of this living matter. The nucleus was looked upon as simply a denser portion of the protoplasm, and the nucleolus as a still denser portion. Now it is believed that the nucleus is a distinct substance, and a permanent organ of the cell. The remaining portion of the protoplasm is now usually spoken of as the cytoplasm.

In spirogyra then the cytoplasm in each cell consists of a layer which lines the inside of the cell wall, a nuclear layer, which surrounds the nucleus, and radiating strands which connect the nucleus and wall layers, thus suspending the nucleus near the center of the cell. But it seems best in this elementary study to use the term protoplasm in its general sense.

Protoplasm in mucor.

12. Let us now examine in a similar way another of the simple plants with the special object in view of demonstrating the protoplasm. For this purpose we may take one of the plants belonging to the group of fungi. These plants possess no chlorophyll. One of several species of mucor, a common mould, is readily obtainable, and very suitable for this study.[3]

13. Mycelium of mucor.—A few days after sowing in some gelatinous culture medium we find slender, hyaline threads, which are very much branched, and, radiating from a central point, form circular colonies, if the plant has not been too thickly sown, as shown in [fig. 6]. These threads of the fungus form the mycelium. From these characters of the plant, which we can readily see without the aid of a microscope, we note how different it is from spirogyra.

To examine for protoplasm let us lift carefully a thin block of gelatine containing the mucor threads, and mount it in water on a glass slip. Under the microscope we see only a small portion of the branched threads. In addition to the absence of chlorophyll, which we have already noted, we see that the mycelium is not divided at short intervals into cells, but appears like a delicate tube with branches, which become successively smaller toward the ends.

14. Appearance of the protoplasm.—Within the tube-like thread now note the protoplasm. It has the same general appearance as that which we noted in spirogyra. It is slimy, or semi-fluid, partly hyaline, and partly granular, the granules consisting of minute particles (the microsomes). While in mucor the protoplasm has the same general appearance as in spirogyra, its arrangement is very different. In the first place it is plainly continuous throughout the tube. We do not see the prominent radiations of strands around a large nucleus, but still the protoplasm does not fill the interior of the threads. Here and there are rounded clear spaces termed vacuoles, which are filled with the watery fluid, cell-sap. The nuclei in mucor are very minute, and cannot be seen except after careful treatment with special reagents.

Fig. 6.
Colonies of mucor.

15. Movement of the protoplasm in mucor.—While examining the protoplasm in mucor we are likely to note streaming movements. Often a current is seen flowing slowly down one side of the thread, and another flowing back on the other side, or it may all stream along in the same direction.

16. Test for protoplasm.—Now let us treat the threads with a solution of iodine. The yellowish-brown color appears which is characteristic of protoplasm when subject to this reagent. If we attempt to stain the living protoplasm with a one per cent aqueous solution of eosin it resists it for a time, but if we first kill the protoplasm with strong alcohol, it reacts quickly to the application of the eosin. If we treat the living threads with glycerine the protoplasm is contracted away from the wall, as we found to be the case with spirogyra. While the color, form and structure of the plant mucor is different from spirogyra, and the arrangement of the protoplasm within the plant is also quite different, the reactions when treated by certain reagents are the same. We are justified then in concluding that the two plants possess in common a substance which we call protoplasm.

Fig. 7.
Thread of mucor, showing protoplasm and vacuoles.

Protoplasm in nitella.

[17.] One of the most interesting plants for the study of one remarkable peculiarity of protoplasm is Nitella. This plant belongs to a small group known as stoneworts. They possess chlorophyll, and, while they are still quite simple as compared with the higher plants, they are much higher in the scale than spirogyra or mucor.

18. Form of nitella.—A common species of nitella is Nitella flexilis. It grows in quiet pools of water. The plant consists of a main axis, in the form of a cylinder. At quite regular intervals are whorls of several smaller thread-like outgrowths, which, because of their position, are termed “leaves,” though they are not true leaves. These are branched in a characteristic fashion at the tip. The main axis also branches, these branches arising in the axil of a whorl, usually singly. The portions of the axis where the whorls arise are the nodes. Each node is made up of a number of small cells definitely arranged. The portion of the axis between two adjacent whorls is an internode. These internodes are peculiar. They consist of but a single “cell,” and are cylindrical, with closed ends. They are sometimes 5-10 cm. long.

Fig. 8.

Portion of plant
nitella.

19. Internode of nitella.—For the study of an internode of nitella, a small one, near the end, or the ends of one of the “leaves” is best suited, since it is more transparent. A small portion of the plant should be placed on the glass slip in water with the cover glass over a tuft of the branches near the growing end. Examined with the microscope the green chlorophyll bodies, which form oval or oblong discs, are seen to be very numerous. They lie quite closely side by side and form in perfect rows along the inner surface of the wall. One peculiar feature of the arrangement of the chlorophyll bodies is that there are two lines, extending from one end of the internode to the other on opposite sides, where the chlorophyll bodies are wanting. These are known as neutral lines. They run parallel with the axis of the internode, or in a more or less spiral manner as shown in [fig. 9].

20. Cyclosis in nitella.—The chlorophyll bodies are stationary on the inner surface of the wall, but if the microscope be properly focussed just beneath this layer we notice a rotary motion of particles in the protoplasm. There are small granules and quite large masses of granular matter which glide slowly along in one direction on a given side of the neutral line. If now we examine the protoplasm on the other side of the neutral line, we see that the movement is in the opposite direction. If we examine this movement at the end of an internode the particles are seen to glide around the end from one side of the neutral line to the other. So that when conditions are favorable, such as temperature, healthy state of the plant, etc., this gliding of the particles or apparent streaming of the protoplasm down one side of the “cell,” and back upon the other, continues in an uninterrupted rotation, or cyclosis. There are many nuclei in an internode of nitella, and they move also.

21. Test for protoplasm.—If we treat the plant with a solution of iodine we get the same reaction as in the case of spirogyra and mucor. The protoplasm becomes yellowish-brown.

Fig. 9.
Cyclosis in nitella.

22. Protoplasm in one of the higher plants.—We now wish to examine, and test for, protoplasm in one of the higher plants. Young or growing parts of any one of various plants—the petioles of young leaves, or young stems of growing plants—are suitable for study. Tissue from the pith of corn (Zea mays) in young shoots just back of the growing point or quite near the joints of older but growing corn stalks furnishes excellent material.

If we should place part of the stem of this plant under the microscope we should find it too opaque for observation of the interior of the cells. This is one striking difference which we note as we pass from the low and simple plants to the higher and more complex ones; not only in general is there an increase of size, but also in general an increase in thickness of the parts. The cells, instead of lying end to end or side by side, are massed together so that the parts are quite opaque. In order to study the interior of the plant we have selected it must be cut into such thin layers that the light will pass readily through them.

For this purpose we section the tissue selected by making with a razor, or other very sharp knife, very thin slices of it. These are mounted in water in the usual way for microscopic study. In this section we notice that the cells are polygonal in form. This is brought about by mutual pressure of all the cells. The granular protoplasm is seen to form a layer just inside the wall, which is connected with the nuclear layer by radiating strands of the same substance. The nucleus does not always lie at the middle of the cell, but often is near one side. If we now apply an alcohol solution of iodine the characteristic yellowish-brown color appears. So we conclude here also that this substance is identical with the living matter in the other very different plants which we have studied.

23. Movement of protoplasm in the higher plants.—Certain parts of the higher plants are suitable objects for the study of the so-called streaming movement of protoplasm, especially the delicate hairs, or thread-like outgrowths, such as the silk of corn, or the delicate staminal hairs of some plants, like those of the common spiderwort, tradescantia, or of the tradescantias grown for ornament in greenhouses and plant conservatories.

Sometimes even in the living cells of the corn plant which we have just studied, slow streaming or gliding movements of the granules are seen along the strands of protoplasm where they radiate from the nucleus. [See note] at close of this chapter.

24. Movement of protoplasm in cells of the staminal hair of “spiderwort.”—A cell of one of these hairs from a stamen of a tradescantia grown in glass houses is shown in [fig. 10]. The nucleus is quite prominent, and its location in the cell varies considerably in different cells and at different times. There is a layer of protoplasm all around the nucleus, and from this the strands of protoplasm extend outward to the wall layer. The large spaces between the strands are, as we have found in other cases, filled with the cell-sap.

Fig. 10.
Cell from stamen hair of tradescantia showing movement of the protoplasm.

An entire stamen, or a portion of the stamen, having several hairs attached, should be carefully mounted in water. Care should be taken that the room be not cold, and if the weather is cold the water in which the preparation is mounted should be warm. With these precautions there should be little difficulty in observing the streaming movement.

The movement is detected by observing the gliding of the granules. These move down one of the strands from the nucleus along the wall layer, and in towards the nucleus in another strand. After a little the direction of the movement in any one portion may be reversed.

25. Cold retards the movement.—While the protoplasm is moving, if we rest the glass slip on a block of ice, the movement will become slower, or will cease altogether. Then if we warm the slip gently, the movement becomes normal again. We may now apply here the usual tests for protoplasm. The result is the same as in the former cases.

26. Protoplasm occurs in the living parts of all plants.—In these plants representing such widely different groups, we find a substance which is essentially alike in all. Though its arrangement in the cell or plant body may differ in the different plants or in different parts of the same plant, its general appearance is the same. Though in the different plants it presents, while alive, varying phenomena, as regards mobility, yet when killed and subjected to well known reagents the reaction is in general identical. Knowing by the experience of various investigators that protoplasm exhibits these reactions under given conditions, we have demonstrated to our satisfaction that we have seen protoplasm in the simple alga, spirogyra, in the common mould, mucor, in the more complex stonewort, nitella, and in the cells of tissues of the highest plants.

27. By this simple process of induction of these facts concerning this substance in these different plants, we have learned an important method in science study. Though these facts and deductions are well known, the repetition of the methods by which they are obtained on the part of each student helps to form habits of scientific carefulness and patience, and trains the mind to logical processes in the search for knowledge.

28. While we have by no means exhausted the study of protoplasm, we can, from this study, draw certain conclusions as to its occurrence and appearance in plants. Protoplasm is found in the living and growing parts of all plants. It is a semi-fluid, or slimy, granular, substance; in some plants, or parts of plants, the protoplasm exhibits a streaming or gliding movement of the granules. It is irritable. In the living condition it resists more or less for some time the absorption of certain coloring substances. The water may be withdrawn by glycerine. The protoplasm may be killed by alcohol. When treated with iodine it becomes a yellowish-brown color.

[Note.] In some plants, like elodea for example, it has been found that the streaming of the protoplasm is often induced by some injury or stimulus, while in the normal condition the protoplasm does not move.

[CHAPTER II.]
ABSORPTION, DIFFUSION, OSMOSE.

29. We may next endeavor to learn how plants absorb water or nutrient substances in solution. There are several very instructive experiments, which can be easily performed, and here again some of the lower plants will be found useful.

30. Osmose in spirogyra.—Let us mount a few threads of this plant in water for microscopic examination, and then draw under the cover glass a five per cent solution of ordinary table salt (NaCl) with the aid of filter paper. We shall soon see that the result is similar to that which was obtained when glycerine was used to extract the water from the cell-sap, and to contract the protoplasmic membrane from the cell wall. But the process goes on evenly and the plant is not injured. The protoplasmic layer contracts slowly from the cell wall, and the movement of the membrane can be watched by looking through the microscope. The membrane contracts in such a way that all the contents of the cell are finally collected into a rounded or oval mass which occupies the center of the cell.

If we now add fresh water and draw off the salt solution, we can see the protoplasmic membrane expand again, or move out in all directions, and occupy its former position against the inner surface of the cell wall. This would indicate that there is some pressure from within while this process of absorption is going on, which causes the membrane to move out against the cell wall.

The salt solution draws water from the cell-sap. There is thus a tendency to form a vacuum in the cell, and the pressure on the outside of the protoplasmic membrane causes it to move toward the center of the cell. When the salt solution is removed and the thread of spirogyra is again bathed with water, the movement of the water is inward in the cell. This would suggest that there is some substance dissolved in the cell-sap which does not readily filter out through the membrane, but draws on the water outside. It is this which produces the pressure from within and crowds the membrane out against the cell wall again.

Fig. 11.

Spirogyra before placing
in salt solution.

Fig. 12.

Spirogyra in
5% salt solution.

Fig. 13.

Spirogyra from salt
solution into water.

31. Turgescence.—Were it not for the resistance which the cell wall offers to the pressure from within, the delicate protoplasmic membrane would stretch to such an extent that it would be ruptured, and the protoplasm therefore would be killed. If we examine the cells at the ends of the threads of spirogyra we shall see in most cases that the cell wall at the free end is arched outward. This is brought about by the pressure from within upon the protoplasmic membrane which itself presses against the cell wall, and causes it to arch outward. This is beautifully shown in the case of threads which are recently broken. The cell wall is therefore elastic; it yields to a certain extent to the pressure from within, but a point is soon reached beyond which it will not stretch, and an equilibrium then exists between the pressure from within on the protoplasmic membrane, and the pressure from without by the elastic cell wall. This state of equilibrium in a cell is turgescence, or such a cell is said to be turgescent, or turgid.

Fig. 14.

Before treatment with
salt solution.

Fig. 15.

After treatment with
salt solution.

Fig. 16.

From salt solution placed
in water.

Figs. 14-16.—Osmosis in threads of mucor.

32. Experiment with beet in salt and sugar solutions.—We may now test the effect of a five per cent salt solution on a portion of the tissues of a beet or carrot. Let us cut several slices of equal size and about 5mm in thickness. Immerse a few slices in water, a few in a five per cent salt solution and a few in a strong sugar solution. It should be first noted that all the slices are quite rigid when an attempt is made to bend them between the fingers. In the course of one or two hours or less, if we examine the slices we shall find that those in water remain, as at first, quite rigid, while those in the salt and sugar solutions are more or less flaccid or limp, and readily bend by pressure between the fingers, the specimens in the salt solution, perhaps, being more flaccid than those in the sugar solution. The salt solution, we judge after our experiment with spirogyra, withdraws some of the water from the cell-sap, the cells thus losing their turgidity and the tissues becoming limp or flaccid from the loss of water.

Fig. 17.

Before treatment with
salt solution.

Fig. 18.

After treatment with
salt solution.

Fig. 19.

From salt solution into
water again.

Figs. 17-19.—Osmosis in cells of Indian corn.

Fig. 20.

Rigid condition of
fresh beet section.

Fig. 21.

Limp condition after
lying in salt
solution.

Fig. 22.

Rigid again after
lying again in water.

Figs. 20-22.—Turgor and osmosis in slices of beet.

33. Let us now remove some of the slices of the beet from the sugar and salt solutions, wash them with water and then immerse them in fresh water. In the course of thirty minutes to one hour, if we examine them again, we find that they have regained, partly or completely, their rigidity. Here again we infer from the former experiment with spirogyra that the substances in the cell-sap now draw water inward; that is, the diffusion current is inward through the cell walls and the protoplasmic membrane, and the tissue becomes turgid again.

Fig. 23.

Before treatment with
salt solution.

Fig. 24.

After treatment with
salt solution.

Fig. 25.

Later stage of
the same.

Figs. 23-25.—Cells from beet treated with salt solution to
show osmosis and movement of the protoplasmic membrane.

34. Osmose in the cells of the beet.—We should now make a section of the fresh tissue of a red colored beet for examination with the microscope, and treat this section with the salt solution. Here we can see that the effect of the salt solution is to draw water out of the cell, so that the protoplasmic membrane can be seen to move inward from the cell wall just as was observed in the case of spirogyra.[4] Now treating the section with water and removing the salt solution, the diffusion current is in the opposite direction, that is inward through the protoplasmic membrane, so that the latter is pressed outward until it comes in contact with the cell wall again, which by its elasticity soon resists the pressure and the cells again become turgid.

35. The coloring matter in the cell-sap does not readily escape from the living protoplasm of the beet.—The red coloring matter, as seen in the section under the microscope, does not escape from the cell-sap through the protoplasmic membrane. When the slices are placed in water, the water is not colored thereby. The same is true when the slices are placed in the salt or sugar solutions. Although water is withdrawn from the cell-sap, this coloring substance does not escape, or if it does it escapes slowly and after a considerable time.

36. The coloring matter escapes from dead protoplasm.—If, however, we heat the water containing a slice of beet up to a point which is sufficient to kill the protoplasm, the red coloring matter in the cell-sap filters out through the protoplasmic membrane and colors the water. If we heat a preparation made for study under the microscope up to the thermal death point we can see here that the red coloring matter escapes through the membrane into the water outside. This teaches that certain substances cannot readily filter through the living membrane of protoplasm, but that they can filter through when the protoplasm is dead. A very important condition, then, for the successful operation of some of the physical processes connected with absorption in plants is that the protoplasm should be in a living condition.

37. Osmose experiments with leaves.—We may next take the leaves of certain plants like the geranium, coleus or other plant, and place them in shallow vessels containing water, salt, and sugar solutions respectively. The leaves should be immersed, but the petioles should project out of the water or solutions. Seedlings of corn or beans, especially the latter, may also be placed in these solutions, so that the leafy ends are immersed. After one or two hours an examination shows that the specimens in the water are still turgid. But if we lift a leaf or a bean plant from the salt or sugar solution, we find that it is flaccid and limp. The blade, or lamina, of the leaf droops as if wilted, though it is still wet. The bean seedling also is flaccid, the succulent stem bending nearly double as the lower part of the stem is held upright. This loss of turgidity is brought about by the loss of water from the tissues, and judging from the experiments on spirogyra and the beet, we conclude that the loss of turgidity is caused by the withdrawal of some of the water from the cell-sap by the strong salt solution.

38. Now if we wash carefully these leaves and seedlings, which have been in the salt and sugar solutions, with water, and then immerse them in fresh water for a few hours, they will regain their turgidity. Here again we are led to infer that the diffusion current is now inward through the protoplasmic membranes of all the living cells of the leaf, and that the resulting turgidity of the individual cells causes the turgidity of the leaf or stem.

Fig. 26.
Seedling of radish,
showing root hairs.

39. Absorption by root hairs.—If we examine seedlings, which have been grown in a germinator or in the folds of paper or cloths so that the roots will be free from particles of soil, we see near the growing point of the roots that the surface is covered with numerous slender, delicate, thread-like bodies, the root hairs. Let us place a portion of a small root containing some of these root hairs in water on a glass slip, and prepare it for examination with the microscope. We see that each thread, or root hair, is a continuous tube, or in other words it is a single cell which has become very much elongated. The protoplasmic membrane lines the wall, and strands of protoplasm extend across at irregular intervals, the interspaces being occupied by the cell-sap.

Fig. 27.
Root hair of corn before and after treatment with 5% salt solution.

We should now draw under the cover glass some of the five per cent salt solution. The protoplasmic membrane moves away from the cell wall at certain points, showing that plasmolysis is taking place, that is, the diffusion current is outward so that the cell-sap loses some of its water, and the pressure from the outside moves the membrane inward. We should not allow the salt solution to work on the root hairs long. It should be very soon removed by drawing in fresh water before the protoplasmic membrane has been broken at intervals, as is apt to be the case by the strong diffusion current and the consequent strong pressure from without. The membrane of protoplasm now moves outward as the diffusion current is inward, and soon regains its former position next the inner side of the cell wall. The root hairs then, like other parts of the plant which we have investigated, have the power of taking up water under pressure.

40. Cell-sap a solution of certain substances.—From these experiments we are led to believe that certain substances reside in the cell-sap of plants, which behave very much like the salt solution when separated from water by the protoplasmic membrane. Let us attempt to interpret these phenomena by recourse to diffusion experiments, where an animal membrane separates two liquids of different concentration.

41. An artificial cell to illustrate turgor.—Fill a small wide-mouthed vial with a very strong sugar solution. Over the mouth tie firmly a piece of bladder membrane. Be certain that as the membrane is tied over the open end of the vial, the sugar solution fills it in order to keep out air bubbles. Sink the vial in a vessel of fresh water and leave it there for twenty-four hours. Remove the vial and note that the membrane is arched outward. Thrust a sharp needle through the membrane when it is arched outward, and quickly pull it out. The liquid spurts out because of the inside pressure.

Fig. 28.
Puncturing a make-believe cell after it has been lying in water.

Fig. 29.
Same as Fig. 28 after needle is removed.

42. Diffusion through an animal membrane.—For this experiment we may use a thistle tube, across the larger end of which should be stretched and tied tightly a piece of a bladder membrane. A strong sugar solution (three parts sugar to one part water) is now placed in the tube so that the bulb is filled and the liquid extends part way in the neck of the tube. This is immersed in water within a wide-mouth bottle, the neck of the tube being supported in a perforated cork in such a way that the sugar solution in the tube is on a level with the water in the bottle or jar. In a short while the liquid begins to rise in the thistle tube, in the course of several hours having risen several centimeters. The diffusion current is thus stronger through the membrane in the direction of the sugar solution, so that this gains more water than it loses.

We have here two liquids separated by an animal membrane, water on the one hand which diffuses readily through the membrane, while on the other is a solution of sugar which diffuses through the animal membrane with difficulty. The water, therefore, not containing any solvent, according to a general law which has been found to obtain in such cases, diffuses more readily through the membrane into the sugar solution, which thus increases in volume, and also becomes more dilute. The bladder membrane is what is sometimes called a diffusion membrane, since the diffusion currents travel through it.

43. In this experiment then the bulk of the sugar solution is increased, and the liquid rises in the tube by this pressure above the level of the water in the jar outside of the thistle tube. The diffusion of liquids through a membrane is osmosis.

44. Importance of these physical processes in plants.—Now if we recur to our experiment with spirogyra we find that exactly the same processes take place. The protoplasmic membrane is the diffusion membrane, through which the diffusion takes place. The salt solution which is first used to bathe the threads of the plant is a stronger solution than that of the cell-sap within the cell. Water therefore is drawn out of the cell-sap, but the substances in solution in the cell-sap do not readily move out. As the bulk of the cell-sap diminishes the pressure from the outside pushes the protoplasmic membrane away from the wall. Now when we remove the salt solution and bathe the thread with water again, the cell-sap, being a solution of certain substances, diffuses with more difficulty than the water, and the diffusion current is inward, while the protoplasmic membrane moves out against the cell wall, and turgidity again results. Also in the experiments with salt and sugar solutions on the leaves of geranium, on the leaves and stems of the seedlings, on the tissues and cells of the beet and carrot, and on the root hairs of the seedlings, the same processes take place.

These experiments not only teach us that in the protoplasmic membrane, the cell wall, and the cell-sap of plants do we have structures which are capable of performing these physical processes, but they also show that these processes are of the utmost importance to the plant; not only in giving the plant the power to take up solutions of nutriment from the soil, but they serve also other purposes, as we shall see later.

[CHAPTER III.]
HOW PLANTS OBTAIN WATER.

In connection with the study of the means of absorption from the soil or water by plants, it will be found convenient to observe carefully the various forms of the plant. Without going into detail here, the suggestion is made that simple thread forms like spirogyra, œdogonium, and vaucheria; expanded masses of cells as are found in the thalloid liverworts, the duckweed, etc., be compared with those liverworts, and with the mosses, where leaf-like expansions of a central axis have been differentiated. We should then note how this differentiation, from the physiological standpoint, has been carried farther in the higher land plants.

45. Absorption by Algæ and Fungi.—In the simpler forms of plant life, as in spirogyra and many of the algæ and fungi, the plant body is not differentiated into parts.[5] In many other cases the only differentiation is between the growing part and the fruiting part. In the algæ and fungi there is no differentiation into stem and leaf, though there is an approach to it in some of the higher forms. Where this simple plant body is flattened, as in the sea-wrack, or ulva, it is a frond. The Latin word for frond is thallus, and this name is applied to the plant body of all the lower plants, the algæ and fungi. The algæ and fungi together are sometimes called the thallophytes, or thallus plants. The word thallus is also sometimes applied to the flattened body of the liverworts. In the foliose liverworts and mosses there is an axis with leaf-like expansions. These are believed by some to represent true stems and leaves, by others to represent a flattened thallus in which the margins are deeply and regularly divided, or in which the expansion has only taken place at regular intervals.

In nearly all of the algæ the plant body is submerged in water. In these cases absorption takes place through all portions of the surface in contact with the water, as in spirogyra, vaucheria, and all of the larger seaweeds. Comparatively few of the algæ grow on the surfaces of rocks or trees. In these examples it is likely that at times only portions of the plant body serve in the process of absorption of water from the substratum. A few of the algæ are parasitic, living in the tissues of higher plants, where they are surrounded by the water or liquids within the host. Absorption takes place in the same way in many of the fungi. The aquatic fungi are immersed in water. In other forms, like mucor, a portion of the mycelium is within the substratum, and being bathed by the water or watery solutions absorbs the same, while the fruiting portion and the aerial mycelium obtain their water and food solutions from the mycelium in the substratum. In higher fungi, like the mushrooms, the mycelium within the ground or decaying wood absorbs the water necessary for the fruiting portion; while in the case of the parasitic fungi the mycelium lies in the water or liquid within the host.

Fig. 30.
Thallus of Riccia lutescens.

46. Absorption by liverworts.—In many of the plants termed liverworts the vegetative part of the plant is a thin, flattened, more or less elongated green body known as a thallus.

Riccia.—One of these, belonging to the genus riccia, is shown in [fig. 30]. Its shape is somewhat like that of a minute ribbon which is forked at intervals in a dichotomous manner, the characteristic kind of branching found in these thalloid liverworts. This riccia (known as R. lutescens) occurs on damp soil; long, slender, hair-like processes grow out from the under surface of the thallus which resemble root hairs and serve the same purpose in the processes of absorption. Another species of riccia (R. crystallina) is shown in [fig. 252]. This plant is quite circular in outline and occurs on muddy flats. Some species float on the water.

47. Marchantia.—One of the larger and coarser liverworts is [figured at 31]. This is a very common liverwort, growing in very damp and muddy places and also along the margins of streams, on the mud or upon the surfaces of rocks which are bathed with the water. This is known as Marchantia polymorpha. If we examine the under surface of the marchantia we see numerous hair-like processes which attach the plant to the soil. Under the microscope we see that some of these are similar to the root hairs of the seedlings which we have been studying, and they serve the purpose of absorption. Since, however, there are no roots on the marchantia plant, these hair-like outgrowths are usually termed here rhizoids. In marchantia they are of two kinds, one kind the simple ones with smooth walls, and the other kind in which the inner surfaces of the walls are roughened by processes which extend inward in the form of irregular tooth-like points. Besides the hairs on the under side of the thallus we note especially near the growing end that there are two rows of leaf-like scales, those at the end of the thallus curving up over the growing end, thus serving to protect the delicate tissues at the growing point.

Fig. 31.
Marchantia plant with cupules and gemmæ; rhizoids below.

Fig. 32.

Portion of plant
of Frullania,
a foliose
liverwort.

Fig. 33.

Portion of same
more highly
magnified, showing
overlapping leaves.

Fig. 34.

Under side,
showing forked
under row of
leaves and lobes
of lateral leaves.

48. Frullania.—In [fig. 32] is shown another liverwort, which differs greatly in form from the ones we have just been studying in that there is a well-defined axis with lateral leaf-like outgrowths. Such liverworts are called foliose liverworts. Besides these two quite prominent rows of leaves there is a third row of poorly developed leaves on the under surface. Also from the under surface of the axis we see here and there slender outgrowths, the rhizoids, through which much of the water is absorbed.

Fig. 35.
Foliose liverwort (bazzania) showing
dichotomous branching and overlapping leaves.

49. Absorption by the mosses.—Among the mosses, which are usually common in moist and shaded situations, examples are abundant which are suitable for the study of the organs of absorption. If we take for example a plant of mnium (M. affine), which is illustrated in [fig. 36], we note that it consists of a slender axis with thin flat, green, leaf-like expansions, Examining with the microscope the lower end of the axis, which is attached to the substratum, there are seen numerous brown-colored threads more or less branched.

Fig. 36.
Female plant (gametophyte)
of a moss (mnium), showing
rhizoids below, and the
tuft of leaves above,
which protect the
archegonia.

50. Absorption by the higher aquatic plants.—Examples of the water plants which are entirely submerged in water are the water-crowfoots, some of the pondweeds, elodea or water-weeds, the tape-grass, vallisneria, etc. In these plants all parts of the body being submerged, they absorb water with which they are in contact. In other aquatic plants, like the water-lilies, some of the pondweeds, the duck-meats, etc., are only partially submerged in the water; the upper surface of the leaf or of the leaf-like expansion being exposed to the air, while the under surface lies in close contact with the water, and the stems and the petioles of the leaves are also immersed in water. In these plants absorption takes place through those parts in contact with the water.

[51. Absorption by the duck-meats.]—These plants are very curious examples of the higher plants.

Lemna.—One of these is illustrated in [fig. 37]. This is the common duckweed, Lemna trisulca. It is very peculiar in form and in its mode of growth. Each one of the lateral leaf-like expansions extends outwards by the elongation of the basal part, which becomes long and slender. Next, two new lateral expansions are formed on these by prolification from near the base, and thus the plant continues to extend. The plant occurs in ponds and ditches and is sometimes very common and abundant. It floats on the surface of the water. While the flattened part of the plant resembles a leaf, it is really the stem, no leaves being present. This expanded green body is usually termed a “frond.” A single rootlet grows out from the under side and is destitute of root hairs. Absorption of water therefore takes place through this rootlet and through the under side of the “frond.”

Fig. 37.
Fronds of the duckweed (Lemna trisculca).

Fig. 38.
Spirodela polyrhiza.

52. Spirodela polyrhiza.—This is a very curious plant, closely related to the lemna and sometimes placed in the same genus. It occurs in similar situations, and is very readily grown in aquaria. It reminds one of a little insect as seen in [fig. 38]. There are several rootlets on the under side of the frond. Absorption of water takes place here in the same way as in lemna.

53. Absorption in wolffia.—Perhaps the most curious of these modified water plants is the little wolffia, which contains the smallest specimens of the flowering plants. Two species of this genus are shown in [figs. 39-41]. The plant body is reduced to nothing but a rounded or oval green body, which represents the stem. No leaves or roots are present. The plants multiply by “prolification,” the new fronds growing out from a depression on the under side of one end. Absorption takes place through the under surface.

[54. Absorption by land plants.]—Water cultures.—In connection with our inquiry as to how land plants obtain their water, it will be convenient to prepare some water cultures to illustrate this and which can also be used later in our study of nutrition ([Chapter IX]).

Fig. 39.
Young frond of wolffia
growing out of older one.

Fig. 40.
Young frond of wolffia
separating from older one.

Fig. 41.
Another species of
wolffia, the two fronds
still connected.

Chemical analysis shows that certain mineral substances are common constituents of plants. By growing plants in different solutions of these various substances it has been possible to determine what ones are necessary constituents of plant food. While the proportion of the mineral elements which enter into the composition of plant food may vary considerably within certain limits, the concentration of the solutions should not exceed certain limits. A very useful solution is one recommended by Sachs, and is as follows:

55. Formula for water cultures:

The calcium phosphate is only partly soluble. The solution which is not in use should be kept in a dark cool place to prevent the growth of minute algæ.

56. Several different plants are useful for experiments in water cultures, as peas, corn, beans, buckwheat, etc. The seeds of these plants may be germinated, after soaking them for several hours in warm water, by placing them between the folds of wet paper on shallow trays, or in the folds of wet cloth. The seeds should not be kept immersed in water after they have imbibed enough to thoroughly soak and swell them. At the same time that the seeds are placed in damp paper or cloth for germination, one lot of the soaked seeds should be planted in good soil and kept under the same temperature conditions, for control. When the plants have germinated one series should be grown in distilled water, which possesses no plant food; another in the nutrient solution, and still another in the nutrient solution to which has been added a few drops of a solution of iron chloride or ferrous sulphate. There would then be four series of cultures which should be carried out with the same kind of seed in each series so that the comparisons can be made on the same species under the different conditions. The series should be numbered and recorded as follows:

• No. 1, soil.
• No. 2, distilled water.
• No. 3, nutrient solution.
• No. 4, nutrient solution with a few drops of iron solution added.

Fig. 42.
Culture cylinder to
show position of corn
seedling (Hansen).

57. Small jars or wide-mouth bottles, or crockery jars, can be used for the water cultures, and the cultures are set up as follows: A cork which will just fit in the mouth of the bottle, or which can be supported by pins, is perforated so that there is room to insert the seedling, with the root projecting below into the liquid. The seed can be fastened in position by inserting a pin through one side, if it is a large one, or in the case of small seeds a cloth of a coarse mesh can be tied over the mouth of the bottle instead of using the cork. After properly setting up the experiments the cultures should be arranged in a suitable place, and observed from time to time during several weeks. In order to obtain more satisfactory results several duplicate series should be set up to guard against the error which might arise from variation in individual plants and from accident. Where there are several students in a class, a single series set up by several will act as checks upon one another. If glass jars are used for the liquid cultures they should be wrapped with black paper or cloth to exclude the light from the liquid, otherwise numerous minute algæ are apt to grow and interfere with the experiment. Or the jars may be sunk in pots of earth to serve the same purpose. If crockery jars are used they will not need covering.

58. For some time all the plants grow equally well, until the nutriment stored in the seed is exhausted. The numbers 1, 3 and 4, in soil and nutrient solutions, should outstrip number 2, the plants in the distilled water. No. 4 in the nutrient solution with iron, having a perfect food, compares favorably with the plants in the soil.

59. Plants take liquid food from the soil.—From these experiments then we judge that such plants take up the food they receive from the soil in the form of a liquid, the elements being in solution in water.

If we recur now to the experiments which were performed with the salt solution in producing plasmolysis in the cells of spirogyra, in the cells of the beet or corn, and in the root hairs of the corn and bean seedlings, and the way in which these cells become turgid again when the salt solution is removed and they are again bathed with water, we shall have an explanation of the way in which plants take up nutrient solutions of food material through their roots.

Fig. 43.
Section of corn root, showing rhizoids
formed from elongated epidermal cells.

60. How food solutions are carried into the plant.—We can see how water and food solutions are carried into the plant, and we must next turn our attention to the way in which these solutions are carried farther into the plant. We should make a section across the root of a seedling in the region of the root hairs and examine it with the aid of a microscope. We here see that the root hairs are formed by the elongation of certain of the surface cells of the root. These cells elongate perpendicularly to the root, and become 3mm to 6mm long. They are flexuous or irregular in outline and cylindrical, as shown in [fig. 43]. The end of the hair next the root fits in between the adjacent superficial cells of the root and joins closely to the next deeper layer of cells. In studying the section of the young root we see that the root is made up of cells which lie closely side by side, each with its wall, its protoplasm and cell-sap, the protoplasmic membrane lying on the inside of each cell wall.

61. In the absorption of the watery solutions of plant food by the root hairs, the cell-sap, being a more concentrated solution, gains some of the former, since the liquid of less concentration flows through the protoplasmic membrane into the more concentrated cell-sap, increasing the bulk of the latter. This makes the root hairs turgid, and at the same time dilutes the cell-sap so that the concentration is not so great. The cells of the root lying inside and close to the base of the root hairs have a cell-sap which is now more concentrated than the diluted cell-sap of the hairs, and consequently gain some of the food solutions from the latter, which tends to lessen the content of the root hairs and also to increase the concentration of the cell-sap of the same. This makes it possible for the root hairs to draw on the soil for more of the food solutions, and thus, by a variation in the concentration of the substances in solution in the cell-sap of the different cells, the food solutions are carried along until they reach the vascular bundles, through which the solutions are carried to distant parts of the plant. Some believe that there is a rhythmic action of the elastic cell walls in these cells between the root hairs and the vascular bundles. This occurs in such a way that, after the cell becomes turgid, it contracts, thus reducing the size of the cell and forcing some of the food solutions into the adjacent cells, when by absorption of more food solutions, or water, the cell increases in turgidity again. This rhythmic action of the cells, if it does take place, would act as a pump to force the solutions along, and would form one of the causes of root pressure.

62. How the root hairs get the watery solutions from the soil.—If we examine the root hairs of a number of seedlings which are growing in the soil under normal conditions, we shall see that a large quantity of soil readily clings to the roots. We should note also that unless the soil has been recently watered there is no free water in it; the soil is only moist. We are curious to know how plants can obtain water from soil which is not wet. If we attempt to wash off the soil from the roots, being careful not to break away the root hairs, we find, that small particles cling so tenaciously to the root hairs that they are not removed. Placing a few such root hairs under the microscope it appears as if here and there the root hairs were glued to the minute soil particles.

Fig. 44.
Root hairs of corn seedling with soil particles adhering closely.

63. If now we take some of the soil which is only moist, weigh it, and then permit it to become quite dry on exposure to dry air, and weigh again, we find that it loses weight in drying. Moisture has been given off. This moisture, it has been found, forms an exceedingly thin film on the surface of the minute soil particles. Where these soil particles lie closely together, as they usually do when massed together in the pot or elsewhere, this thin film of moisture is continuous from the surface of one particle to that of another. Thus the soil particles which are so closely attached to the root hairs connect the surface of the root hairs with this film of moisture. As the cell-sap of the root hairs draws on the moisture film with which they are in contact, the tension of this film is sufficient to draw moisture from distant particles. In this way the roots are supplied with water in soil which is only moist.

64. Plants cannot remove all the moisture from the soil.—If we now take a potted plant, or a pot containing a number of seedlings, place it in a moderately dry room, and do not add water to the soil we find in a few days that the plant is wilting. The soil if examined will appear quite dry to the sense of touch. Let us weigh some of this soil, then dry it by artificial heat, and weigh again. It has lost in weight. This has been brought about by driving off the moisture which still remained in the soil after the plant began to wilt. This teaches that while plants can obtain water from soil which is only moist or which is even rather dry, they are not able to withdraw all the moisture from the soil.

Fig. 45.
Experiment to show
root pressure
(Detmer).

65. “Root pressure” or exudation pressure.—It is a very common thing to note, when certain shrubs or vines are pruned in the spring, the exudation of a watery fluid from the cut surfaces. In the case of the grape vine this has been known to continue for a number of days, and in some cases the amount of liquid, called “sap,” which escapes is considerable. In many cases it is directly traceable to the activity of the roots, or root hairs, in the absorption of water from the soil. For this reason the term root pressure has been used to denote the force exerted in supplying the water from the soil. But there are some who object to the use of this term “root pressure.” The principal objection is that the pressure which brings about the phenomenon known as “bleeding” by plants is not present in the roots alone. This pressure exists under certain conditions in all parts of the plant. The term exudation pressure has been proposed in lieu of root pressure. It should be remembered that the movement of water in the plant is started by the pressure which exists in the root. If the term “root pressure” is used, it should be borne clearly in mind that it does not express the phenomenon exactly in all cases.

Root pressure may be measured.—It is possible to measure not only the amount of water which the roots will raise in a given time, but also to measure the force exerted by the roots during root pressure. It has been found that root pressure in the case of the nettle is sufficient to hold a column of water about 4.5 meters (15 ft.) high (Vines), while the root pressure of the vine (Hales, 1721) will hold a column of water about 10 meters (36.5 ft.) high, and the birch (Betula lutea) (Clark, 1873) has a root pressure sufficient to hold a column of water about 25 meters (84.7 ft.) high.

66. Experiment to demonstrate root pressure.—By a very simple method this lifting of water by root pressure is shown. During the summer season plants in the open may be used if it is preferred, but plants grown in pots are also very serviceable, and one may use a potted begonia or balsam, the latter being especially useful. The plants are usually convenient to obtain from the greenhouses, to illustrate this phenomenon. The stem is cut off rather close to the soil and a long glass tube is attached to the cut end of the stem, still connected with the roots, by the use of rubber tubing, as shown in [figure 45], and a very small quantity of water may be poured in to moisten the cut end of the stem. In a few minutes the water begins to rise in the glass tube. In some cases it rises quite rapidly, so that the column of water can readily be seen to extend higher and higher up in the tube when observed at quite short intervals. (To measure the force of root pressure is rather difficult for elementary work. To measure it see Ganong, Plant Physiology, pp. 67, 68, or some other book for advanced work.)

67. In either case where the experiment is continued for several days it is noticed that the column of water or of mercury rises and falls at different times during the same day, that is, the column stands at varying heights; or in other words the root pressure varies during the day. With some plants it has been found that the pressure is greatest at certain times of the day, or at certain seasons of the year. Such variation of root pressure exhibits what is termed a periodicity, and in the case of some plants there is a daily periodicity; while in others there is in addition an annual periodicity. With the grape vine the root pressure is greatest in the forenoon, and decreases from 12-6 p.m., while with the sunflower it is greatest before 10 a.m., when it begins to decrease. Temperature of the soil is one of the most important external conditions affecting the activity of root pressure.

[CHAPTER IV.]
TRANSPIRATION, OR THE LOSS OF WATER
BY PLANTS.

68. We should now inquire if all the water which is taken up in excess of that which actually suffices for turgidity is used in the elaboration of new materials of construction. We notice when a leaf or shoot is cut away from a plant, unless it is kept in quite a moist condition, or in a damp, cool place, that it becomes flaccid, and droops. It wilts, as we say. The leaves and shoot lose their turgidity. This fact suggests that there has been a loss of water from the shoot or leaf. It can be readily seen that this loss is not in the form of drops of water which issue from the cut end of the shoot or petiole. What then becomes of the water in the cut leaf or shoot?

69. Loss of water from excised leaves.—Let us take a handful of fresh, green, rather succulent leaves, which are free from water on the surface, and place them under a glass bell jar, which is tightly closed below but which contains no water. Now place this in a brightly lighted window, or in sunlight. In the course of fifteen to thirty minutes we notice that a thin film of moisture is accumulating on the inner surface of the glass jar. After an hour or more the moisture has accumulated so that it appears in the form of small drops of condensed water. We should set up at the same time a bell jar in exactly the same way but which contains no leaves. In this jar there is no condensed moisture on the inner surface. We thus are justified in concluding that the moisture in the former jar comes from the leaves. Since there is no visible water on the surfaces of the leaves, or at the cut ends, before it may have condensed there, we infer that the water escapes from the leaves in the form of water vapor, and that this water vapor, when it comes in contact with the surface of the cold glass, condenses and forms the moisture film, and later the drops of water. The leaves of these cut shoots therefore lose water in the form of water vapor, and thus a loss of turgidity results.

Fig. 46.
To show loss of water from leaves, the leaves just covered.

Fig. 47.
After a few hours drops of water have accumulated
on the inside of the jar covering the leaves.

70. Loss of water from growing plants.—Suppose we now take a small and actively growing plant in a pot, and cover the pot and the soil with a sheet of rubber cloth or flexible oilcloth which fits tightly around the stem of the plant so that the moisture from the soil or from the surface of the pot cannot escape. Then place a bell jar over the plant, and set in a brightly lighted place, at a temperature suitable for growth. In the course of a few minutes on a dry day a moisture film forms on the inner surface of the glass, just as it did in the case of the glass jar containing the cut shoots and leaves. Later the moisture has condensed so that it is in the form of drops. If we have the same leaf surface here as we had with the cut shoots, we shall probably find that a larger amount of water accumulates on the surface of the jar from the plant that is still attached to its roots.

71. Water escapes from the surfaces of living leaves in the form of water vapor.—This living plant then has lost water, which also escapes in the form of water vapor. Since here there are no cut places on the shoots or leaves, we infer that the loss of water vapor takes place from the surfaces of the leaves and from the shoots. It is also to be noted that, while this plant is losing water from the surfaces of the leaves, it does not wilt or lose its turgidity. The roots by their activity and pressure supply water to take the place of that which is given off in the form of water vapor. This loss of water in the form of water vapor by plants is transpiration.

72. A test for the escape of water vapor from plants.—Make a solution of cobalt chloride in water. Saturate several pieces of filter paper with it. Allow them to dry. The water solution of cobalt chloride is red. The paper is also red when it is moist, but when it is thoroughly dry it is blue. It is very sensitive to moisture and the moisture of the air is often sufficient to redden it. Before using dry the paper in an oven or over a flame.

73. Take two bell jars, as shown in [fig. 49]. Under one place a potted plant, the pot and earth being covered by oiled paper. Or cover the plant with a fruit jar. To a stake in the pot pin a piece of the dried cobalt paper, and at the same time pin to a stake, in another jar covering no plant, another piece of cobalt paper. They should both be put under the jars at the same time. In a few moments the paper in the jar with the plant will begin to redden. In a short while, ten or fifteen minutes, probably, it will be entirely red, while the paper under the other jar will remain blue, or be only slightly reddened. The water vapor passing off from the living plant comes in contact with the sensitive cobalt chloride in the paper and reddens it before there is sufficient vapor present to condense as a film of moisture on the surface of the jar.

Fig. 48.

Fig. 49.

Fig. 48.—Water vapor is given off by the leaves when attached to the living plant. It condenses into drops of water on the cool surface of the glass covering the plant.

Fig. 49.—A good way to show that the water passes off from the leaves in the form of water vapor.

74. Experiment to compare loss of water in a dry and a humid atmosphere.—We should now compare the escape of water from the leaves of a plant covered by a bell jar, as in the last experiment, with that which takes place when the plant is exposed in a normal way in the air of the room or in the open. To do this we should select two plants of the same kind growing in pots, and of approximately the same leaf surface. The potted plants are placed one each on the arms of a scale. One of the plants is covered in this position with a bell jar. With weights placed on the pan of the other arm the two sides are balanced. In the course of an hour, if the air of the room is dry, moisture has probably accumulated on the inner surface of the glass jar which is used to cover one of the plants. This indicates that there has here been a loss of water. But there is no escape of water vapor into the surrounding air so that the weight on this arm is practically the same as at the beginning of the experiment. We see, however, that the other arm of the balance has risen. We infer that this is the result of the loss of water vapor from the plant on that arm. Now let us remove the bell jar from the other plant, and with a cloth wipe off all the moisture from the inner surface, and replace the jar over the plant. We note that the end of the scale which holds this plant is still lower than the other end.

75. The loss of water is greater in a dry than in a humid atmosphere.—This teaches us that while water vapor escaped from the plant under the bell jar, the air in this receiver soon became saturated with the moisture, and thus the farther escape of moisture from the leaves was checked. It also teaches us another very important fact, viz., that plants lose water more rapidly through their leaves in a dry air than in a humid or moist atmosphere. We can now understand why it is that during the very hot and dry part of certain days plants often wilt, while at nightfall, when the atmosphere is more humid, they revive. They lose more water through their leaves during the dry part of the day, other things being equal, than at other times.

76. How transpiration takes place.—Since the water of transpiration passes off in the form of water vapor we are led to inquire if this process is simply evaporation of water through the surface of the leaves, or whether it is controlled to any appreciable extent by any condition of the living plant. An experiment which is instructive in this respect we shall find in a comparison between the transpiration of water from the leaves of a cut shoot, allowed to lie unprotected in a dry room, and a similar cut shoot the leaves of which have been killed.

77. Almost any plant will answer for the experiment. For this purpose I have used the following method. Small branches of the locust (Robinia pseudacacia), of sweet clover (Melilotus alba), and of a heliopsis were selected. One set of the shoots was immersed for a moment in hot water near the boiling point to kill them. The other set was immersed for the same length of time in cold water, so that the surfaces of the leaves might be well wetted, and thus the two sets of leaves at the beginning of the experiment would be similar, so far as the amount of water on their surfaces is concerned. All the shoots were then spread out on a table in a dry room, the leaves of the killed shoots being separated where they are inclined to cling together. In a short while all the water has evaporated from the surface of the living leaves, while the leaves of the dead shoots are still wet on the surface. In six hours the leaves of the dead shoots from which the surface water had now evaporated were beginning to dry up, while the leaves of the living plants were only becoming flaccid. In twenty-four hours the leaves of the dead shoots were crisp and brittle, while those of the living shoots were only wilted. In twenty-four hours more the leaves of the sweet clover and of the heliopsis were still soft and flexible, showing that they still contained more water than the killed shoots which had been crisp for more than a day.

78. It must be then that during what is termed transpiration the living plant is capable of holding back the water to some extent, which in a dead plant would escape more rapidly by evaporation. It is also known that a body of water with a surface equal to that of a given leaf surface of a plant loses more water by evaporation during the same length of time than the plant loses by transpiration.

79. Structure of a leaf.—We are now led to inquire why it is that a living leaf loses water less rapidly than dead ones, and why less water escapes from a given leaf surface than from an equal surface of water. To understand this it will be necessary to examine the minute structure of a leaf. For this purpose we may select the leaf of an ivy, though many other leaves will answer equally well. From a portion of the leaf we should make very thin cross-sections with a razor or other sharp instrument. These sections should be perpendicular to the surface of the leaf and should be then mounted in water for microscopic examination.[6]

80. Epidermis of the leaf.—In this section we see that the green part of the leaf is bordered on what are its upper and lower surfaces by a row of cells which possess no green color. The walls of the cells of each row have nearly parallel sides, and the cross walls are perpendicular. These cells form a single layer over both surfaces of the leaf and are termed the epidermis. Their walls are quite stout and the outer walls are cuticularized.

Fig. 50.
Section through ivy leaf showing
communication between stomate
and the large intercellular spaces
of the leaf, stoma closed.

Fig. 51.
Stoma open.

Fig. 52.
Stoma closed.

Figs. 51, 52.—Section through stomata of ivy leaf.

81. Soft tissue of the leaf.—The cells which contain the green chlorophyll bodies are arranged in two different ways. Those on the upper side of the leaf are usually long and prismatic in form and lie closely parallel to each other. Because of this arrangement of these cells they are termed the palisade cells, and form what is called the palisade layer. The other green cells, lying below, vary greatly in size in different plants and to some extent also in the same plant. Here we notice that they are elongated, or oval, or somewhat irregular in form. The most striking peculiarity, however, in their arrangement is that they are not usually packed closely together, but each cell touches the other adjacent cells only at certain points. This arrangement of these cells forms quite large spaces between them, the intercellular spaces. If we should examine such a section of a leaf before it is mounted in water we would see that the intercellular spaces are not filled with water or cell-sap, but are filled with air or some gas. Within the cells, on the other hand, we find the cell-sap and the protoplasm.

82. Stomata.—If we examine carefully the row of epidermal cells on the under surface of the leaf, we find here and there a peculiar arrangement of cells shown at figs. [51], [52]. This opening through the epidermal layer is a stoma. The cells which immediately surround the openings are the guard cells. The form of the guard cells can be better seen if we tear a leaf in such a way as to strip off a short piece of the lower epidermis, and mount this in water. The guard cells are nearly crescent-shaped, and the stoma is elliptical in outline. The epidermal cells are very irregular in outline in this view. We should also note that while the epidermal cells contain no chlorophyll, the guard cells do.

Fig. 53.
Portion of epidermis of ivy, showing irregular
epidermal cells, stoma and guard cells.

82a. In the ivy leaf the guard cells are quite plain, but in most plants the form as seen in cross-section is irregular in outline, as shown in [fig. 53a], which is from a section of a wintergreen leaf. This leaf is interesting because it shows the characteristic structure of leaves of many plants growing in soil where absorption of water by the roots is difficult owing to the cold water, acids, or salts in the water or soil, or in dry soil (see Chapters [47], 54, 55). The cuticle over the upper epidermis is quite thick. This lessens the loss of water by the leaf. The compact palisades of cells are in two to three cell layers, also reducing the loss of water.