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.

=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.

=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_.

=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.

=82=_a_. 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. 53_a_, 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.

=83. The living protoplasm retards the evaporation of water from the leaf.=—If we now take into consideration a few facts which we have learned in a previous chapter, with reference to the physical properties of the living cell, we shall be able to give a partial explanation of the comparative slowness with which the water escapes from the leaves. The inner surfaces of the cell walls are lined with the membrane of protoplasm, and within this is the cell-sap. These cells have become turgid by the absorption of the water which has passed up to them from the roots. While the protoplasmic membrane of the cells does not readily permit the water to filter through, yet it is saturated with water, and the elastic cell wall with which it is in contact is also saturated. From the cell wall the water evaporates into the intercellular spaces. But the water is given up slowly through the protoplasmic membrane, so that the water vapor cannot be given off as rapidly from the cell walls as it could if the protoplasm were dead. The living protoplasmic membrane then which is only slowly permeable to the water of the cell-sap is here a very important factor in checking the too rapid loss of water from the leaves.

By an examination of our leaf section we see that the intercellular spaces are all connected, and that the stomata, where they occur, open also into intercellular spaces. There is here an opportunity for the water vapor in the intercellular spaces to escape when the stomata are open.

=84. Action of the stomata.=—The guard cells serve an important function in regulating transpiration. During normal transpiration the guard cells are turgid and their peculiar form then causes them to arch away from each other, allowing the escape of water vapor. When the air becomes too dry transpiration is in excess of absorption by the roots. The guard cells lose some of their water, and collapse so that their inner faces meet in a straight line and close the stoma. Thus the rapid transpiration is checked. Some evaporation of water vapor, however, takes place through the epidermal cells, and if the air remains too dry, the leaves eventually become flaccid and droop. During the day the effect of sunlight is to increase certain sugars or salts in the guard cells so that they readily become turgid and open the stomates, but at night the cell-sap is less concentrated and the stomates are usually closed. Light therefore favors transpiration, while in darkness transpiration is checked.

=85. Compare transpiration from the two surfaces of the leaf.=—This can be done by using the cobalt chloride paper. This paper can be kept from year to year and used repeatedly. It is thus a very simple matter to make these experiments. Provide two pieces of glass (discarded glass negatives, cleaned, are excellent), two pieces of cobalt chloride paper, and some geranium leaves entirely free from surface water. Dry the paper until it is blue. Place one piece of the paper on a glass plate; place the geranium leaf with the under side on the paper. On the upper side of the leaf now place the other cobalt paper, and next the second piece of glass. On the pile place a light weight to keep the parts well in contact. In fifteen or twenty minutes open and examine. The paper next the under side of the geranium leaf is red where it lies under the leaf. The paper on the upper side is only slightly reddened. The greater loss of water, then, is through the under side of the geranium leaf. This is true of a great many leaves, but it is not true of all.

=86. Negative pressure.=—This is not only indicated by the drooping of the leaves, but may be determined in another way. If the shoot of such a plant be cut underneath mercury, or underneath a strong solution of eosin, it will be found that some of the mercury or eosin, as the case may be, will be forcibly drawn up into the stem toward the roots. This is seen on quickly splitting the cut end of the stem. When plants in the open cannot be obtained in this condition, one may take a plant like a balsam plant from the greenhouse, or some other potted plant, knock it out of the pot, free the roots from the soil and allow to partly wilt. The stem may then be held under the eosin solution and cut.

=87. Lifting power of transpiration.=—Not only does transpiration go on quite independently of root pressure, as we have discovered from other experiments, but transpiration is capable of exerting a lifting power on the water in the plant. This may be demonstrated in the following way: Place the cut end of a leafy shoot in one end of a U tube and fit it water-tight. Partly fill this arm of the U tube with water, and add mercury to the other arm until it stands at a level in the two arms as in fig. 54. In a short time we note that the mercury is rising in the tube.

=88. Root pressure may exceed transpiration.=—If we cover small actively growing plants, such as the pea, corn, wheat, bean, etc., with a bell jar, and place them in the sunlight where the temperature is suitable for growth, in a few hours, if conditions are favorable, we shall see that there are drops of water standing out on the margins of the leaves. These drops of water have exuded through the ordinary stomata, or in other cases what are called water stomata, through the influence of root pressure. The plant being covered by the glass jar, the air soon becomes saturated with moisture and transpiration is checked. Root pressure still goes on, however, and the result is shown in the exuding drops. Root pressure is here in excess of transpiration. This phenomenon is often to be observed during the summer season in the case of low-growing plants. During the bright warm day transpiration equals, or may be in excess of, root pressure, and the leaves are consequently flaccid. As nightfall comes on the air becomes more moist, and the conditions of light are such also that transpiration is lessened. Root pressure, however, is still active because the soil is still warm. In these cases drops of water may be seen exuding from the margins of the leaves due to the excess of root pressure over transpiration. Were it not for this provision for the escape of the excess of water raised by root pressure, serious injury by lesions, as a result of the great pressure, might result. The plant is thus to some extent a self-regulatory piece of apparatus so far as root pressure and transpiration are concerned.

=89. Injuries caused by excessive root pressure.=—Some varieties of tomatoes when grown in poorly lighted and poorly ventilated greenhouses suffer serious injury through lesions of the tissues. This is brought about by the cells at certain parts becoming charged so full with water through the activity of root pressure and lessened transpiration, assisted also probably by an accumulation of certain acids in the cell-sap which cannot be got rid of by transpiration. Under these conditions some of the cells here swell out, forming extensive cushions, and the cell walls become so weakened that they burst. It is possible to imitate the excess of root pressure in the case of some plants by connecting the stems with a system of water pressure, when very quickly the drops of water will begin to exude from the margins of the leaves.

=90.= It should be stated that in reality there is no difference between transpiration and evaporation, if we bear in mind that evaporation takes place more slowly from living plants than from dead ones, or from an equal surface of water.

=91.= The escape of water vapor is not the only function of the stomata. The exchange of gases takes place through them as we shall later see. A large number of experiments show that normally the stomata are open when the leaves are turgid. But when plants lose excessive quantities of water on dry and hot days, so that the leaves become flaccid, the guard cells automatically close the stomata to check the escape of water vapor. Some water escapes through the epidermis of many plants, though the cuticularized membrane of the epidermis largely prevents evaporation. In arid regions plants are usually provided with an epidermis of several layers of cells to more securely prevent evaporation there. In such cases the guard cells are often protected by being sunk deeply in the epidermal layer.

=92. Demonstration of stomates and intercellular air spaces.=—A good demonstration of the presence of stomates in leaves, as well as the presence and intercommunication of the intercellular spaces, can be made by blowing into the cut end of the petiole of the leaf of a calla lily, the lamina being immersed in water. The air is forced out through the stomata and rises as bubbles to the surface of the water. At the close of the experiment some of the air bubbles will still be in contact with the leaf surface at the opening of the stomata. The pressure of the water gradually forces this back into the leaf. Other plants will answer for the experiment, but some are more suitable than others.

=92a. Number of stomata.=—The larger number of stomata are on the under side of the leaf. (In leaves which float on the surface of the water all of the stomata are on the upper side of the leaf, as in the water-lily.) It has been estimated by investigation that in general there are 40-300 stomata to the square millimeter of surface. In some plants this number is exceeded, as in the olive, where there are 625. In an entire leaf of Brassica rapa there are about 11,000,000 stomata, and in an entire leaf of the sunflower there are about 13,000,000 stomata.

=92b. Amount of water transpired by plants.=—The amount of water transpired by plants is very great. According to careful estimates a sunflower 6 feet high transpires on the average about one quart per day; an acre of cabbages 2,000,000 quarts in four months; an oak tree with 700,000 leaves transpires about 180 gallons of water per day. According to von Höhnel, a beech tree 110 years old transpired about 2250 gallons of water in one summer. A hectare of such trees (about 400 on 2½ acres) would at the same rate transpire about 900,000 gallons, or about 30,000 barrels in one summer.

FOOTNOTE:

[6] Demonstrations may be made with prepared sections of leaves,