The plant a machine into which energy and material are taken - Carbon assimilation - Feeding - Accumulation and transformations in the plant. The action of light - The chlorophyll-function.

The relations of the plant to the environment can only be understood by taking into account the results of modern physiological discoveries. These teach us that the living plant is a highly complex machine, the details of its organisation and structure being much more numerous and much more closely correlated at numerous points, than the parts of any other machine known to us.

They also teach us that it is supplied with energy from without, as any other machine; and that when so supplied, and properly working, the living structure or machinery does work, also as other machines. But modern physiology goes further, in that it renders some account of the ways by which the external energy is taken into the plant, and there applied to do work, or stored up for a time in order that it may be used to do work at some future time.

The accumulation of energy thus ensured is associated with corresponding changes of material substance, and the principal means for bringing this about is recognised in the assimilation of carbon-dioxide - photo-synthesis.

In this process energy enters the chlorophyll-corpuscle in the form of the radiant energy of the sun, it is there directed in the mechanism of the protoplasm, so as to do work on the molecules of water and carbon-dioxide which have also been brought into the machinery; this it does, breaking asunder their stable structure into unstable bodies, which then re-combine in different ways to form a carbohydrate, such as starch, and this starch is temporarily stored as grains, while oxygen escapes.

Each starch-grain, therefore, is to be regarded as a packet of matter and of potential energy, as it were, capable of yielding up the latter at any future time, when put under such circumstances that it must do so. Such stores of energy-yielding substance, if I may use the much-abused phrase, form the principal food of the plant - or of an animal, if it steps in and takes them - and we now see that the process of carbon-dioxide assimilation, as it has perhaps unfortunately been called, is not the same thing as the process of feeding, for the feeding - i.e. the nutrition proper - of the plant does not begin until the food has been thus obtained.

We now see what the real position of the plant is, to its environment, whether the latter be living or dead. From our point of view, the plant serves as a centre for bringing together the substances obtainable from the soil, and those derived from the atmosphere, and so focussing and directing the radiant energy of the sun upon these substances, that they are broken up, and some of their constituents synthesized, with absorption of energy, into a body, such as starch, containing more energy than did the original substances taken together or separate. It matters little whether the actual carbohydrate thus synthesized is starch, or sugar or inulin: the point is that energy has been gained from outside and bound up with the acquired material for further use. But modern physiology has carried matters much further than this, and especially in the three following directions.

In the first place, it has shown that much of the energy thus stored from without in the plant is again liberated in the process of oxygen respiration, and expended partly as appreciable heat and partly as driving force for stimulating the machinery of the living plant to further activities.

In the second place, part of it is rearranged with the rearrangement of the molecules with which the energy is bound up, as it were, so that work of various kinds is done in the machinery of the plant: I refer to various metabolic and surface-actions resulting from the peculiar mode of presentment of the resulting substances, for instance the production of osmotic pressures in the cell.

And, thirdly, part of the synthesized substance is worked up into higher bodies, by processes which obviously entail the further doing of work on the constituents.

The further pursuit of this theme would evidently carry us beyond the more immediate subject of this book; but I want to make clear that recent researches render it more and more certain that the living plant is a complex piece of co-ordinated machinery which brings together matter and energy from the external universe, and then gets work out of these.

This proposition is the more important because the whole question of the enrichment of our planet with new food, new building materials, and new fuel, to compensate the daily losses, depends on it, and is of course to be referred fundamentally to the acquirement of new supplies of energy from the sun. Enormous activity has been displayed by physiologists, since 1860, in attempting to solve the question, which of the many different rays known to proceed from the sun are absorbed by the chlorophyll-corpuscle, and directed to the performance of the work above referred to.

The names of Draper, Sachs and Pfefifer stand forth prominently as pioneers in this; while those of Lommel, Engelmann, Timiriazeff and Langley have been among the most active in making important contributions to the subject, and in attempting to answer the further questions con nected with the mode in which the chlorophyll is concerned in utilising the energy of the solar radiations. The point is one of supreme importance, because it goes on all fours with modern questions as to the rays of light absorbed or dispersed in our atmosphere at different seasons of the year, or in special climatic conditions, to say nothing of its other scientific aspects. Unfortunately, however, we have no satisfactory explanation of the actual role played by the chlorophyll substance itself, in spite of much industrious work which has been done in the subject in this country and elsewhere. As regards the rays employed, it was first proved that the most effective belong to the red end of the visible spectrum, and that the effect as measured by the amounts of oxygen given off, and of starch formed in given periods of time, is more or less proportionable to the intensity of the solar light. Then it was established that no monochromatic light is so powerful as the white light from which it was obtained, though the relative numbers expressing the activity in the red and yellow regions may stand to those in the blue as something like 12:1. The latest results place the maximum assimilation in the red-orange, and this coincides with the maximum absorption in the chlorophyll. If we may accept the current views as to the distribution of energy in the spectrum of solar light, which depends on the complete absorption of all the rays by a black body, where they are estimated as heat, we have the interesting result that the agricultural or forest plant is adapted to catch and retain, broadly speaking, just those particular rays which possess most energy.

The probability is increasing that the protoplasmic machinery is the really effective mechanism in the process, and we may figure this machinery as so holding or presenting the molecules of carbon-dioxide and water to the impact of the light-vibrations, that the latter are enabled to undo the molecular structure; the atomic combinations thereby liberated may then be supposed to form a body like formic-aldehyde, which by polymerisation becomes a carbohydrate of the nature of a sugar such as glucose, which the protoplasm then builds up into its substance and subsequently deposits as starch, and stores temporarily in the form of grains or as amorphous material.

This is partly hypothetical, and is largely due to the careful deductions of the chemists, but there are very many facts now to hand which bear out its probability, especially the recent advances in our knowledge of the sugars, and the experimental feeding of leaves and plants deprived of starch with such substances as dextrose, levulose, maltose, and other sugars, as well as glycerine and other bodies which should be convertible into, or yield them, if the theory is true. In this last connection, the careful and extensive experiments of Acton, A. Meyer, Boehm, and Laurent should be mentioned. It would be interesting to enlarge upon Engelmann's beautiful physiological experi ments in connection with this subject of absorption of solar energy, where the maximum accumulation of oxygen-loving bacteria at those parts of a green alga which lie in the red-orange of the spectrum, are used as indicators of the maximum oxygen evolution (and therefore of the maximum carbon-dioxide assimilation), but space will not admit of this. For a similar reason I must also pass over the same observer's experiments with plants which assimilate in protoplasm behind a red instead of a green substance, and which absorb chiefly other rays between the yellow and blue, with the remark that they also seem to imply that it is the protoplasmic machinery which turns the energy on to the carbon-dioxide molecule, the coloured screen being secondary in the matter. Recent experiments which show that green plants will not assimilate carbon-dioxide in a light which has passed through a solution of chlorophyll-and therefore left its red rays behind; nor behind a screen of iodine dissolved in carbon-dioxide - which lets no visible rays between the red and blue pass - should be noticed, as showing the importance of the chlorophyll and the special rays referred to, however; and I ought at least to mention Timiriazeff's beautiful proof, published in 1890, that if, on the leaf of a plant left in the dark long enough to render it free of starch, a bright solar spectrum is steadily projected for 3 - 6 hours, the chlorophyll then removed by alcohol and the decolorised leaf placed in iodine, the image of the spectrum is reproduced by the different intensities of the starch bands, blue with iodine, in the different parts. Here, again, the maximum coloration coincides with the maximum absorption in and near the red.

Microscopic observations and photo-chemical experiments alike convince us that the chlorophyll-corpuscle is itself a complex piece of protoplasmic machinery, working for and with the rest of the plant, and there can be little question as to the greater accuracy of our reasoning on the whole question I am discussing, since Meyer, Schimper, Pringsheim, and others have established the importance of its structural peculiarities.

I must now pass on to consider another aspect of the question of carbon-assimilation.

Notes to Chapter 3

In addition to the references in the last chapter, the reader may be referred to Sachs' Lectures, XXV., and Pfeffer's Physiology, pp. 329-356, where the voluminous literature is given.