Kitobni o'qish: «Disease in Plants»
PREFACE
It has often been represented to me that the cultivators of plants, among whom are to be included planters and foresters, as well as agriculturists and gardeners of every kind, are more particularly concerned with, and interested in, the maladies themselves of the plants they grow, than in the life-history of the fungi, insects or other organisms to which they are due, or in the physiological processes which are involved; and although it is impossible to really understand any disease unless we also understand the processes by which it is brought about, there is room for sympathy with the point of view of the cultivator. He says, in effect, "I do not want to know all about the biology of the fungus of wheat-rust, or of the phylloxera, nor do I want to learn what experts can tell me about the action of bacteria in soil, or the process of starch-formation in the leaves: I have neither the time nor the means to master these details. What I want is guidance as to what is wrong with my tomatoes, apple trees, chrysanthemums, fir trees, turnips, etc., and what I am to do to set things right." Just so. With the latter part of this cry one must sympathize, much as a doctor does with the wail of the parent who calls him in to cure his sick child—we need not stop to classify or compare the motives of the parent and the cultivator, and perhaps I had done better to select a breeder of sheep with his flock and a veterinary doctor in the illustration, but we will let it pass; and as regards the former part of the cry, I do not know that the plant-doctor can expect the cultivator to be initiated in the aetiology of the disease any more than the physician expects the parent to understand the biology of the typhoid bacillus. That both the cultivator and the parent would be the better for a real knowledge of the disease in either case must be admitted—nay insisted on, provided the knowledge is real—but we have to deal with facts, and it is a fact that the clients of both doctors are impatient of the details of the case.
Now, of course, I am aware that no short cut or "royal road" to science exists, and if a man is going to train up trees or other plants, he ought to know all about them in health and in sickness, in youth and in old age, and he ought to learn everything about the soil they grow in, the air that surrounds them, the enemies that beset them, and all the multifarious relations of these one to another; but when I look at my boy and reflect how much his nurse, his schoolmaster, his tutor, his doctor, and his parents ought to know successively and simultaneously about him in sickness and in health, and about his surroundings, etc., I begin to wonder whether there is not after all something to be said for the cultivator's point of view.
Moreover, the cultivator knows a good deal about his plants which I do not know, and although I should much like to know it, his plea of want of time rings in my ears and the conviction strikes home that one ought to try and meet his views, and tell him something about disease as manifested in plants without insisting on his becoming a professional mycologist, entomologist, agricultural chemist, and philosopher.
Of course, beyond a certain point, it is his lookout how much the information is worth, and its educational value—a very different matter—is sure to suffer from any restrictions imposed on the treatment of the subject; but if the theme of disease in plants, treated from a general point of view—I was about to write "treated in a popular manner," but that is impossible until physiology and mycology are more widely taught—enables him to understand better the questions he puts to himself, and, still more, if it stimulates him to enquire further into the inexhaustible field of science glimpsed at, something may come of it.
The purpose of these essays is to treat the subject of disease in plants with special reference to the patient itself, and to describe the symptoms it exhibits and the course of the malady, with only such references to the agents which induce or cause disease as are necessary to an intelligent understanding of the subject, and of the kind of treatment called for. Consequently I have avoided any unnecessary classification or elaborate descriptions of parasitic fungi or insects, histological details of the tissues of plants, chemical and physical details regarding the soil, and even matters purely physiological as far as possible. Several admirable works on these subjects are already available, and must be referred to for further details.
It is, however, quite out of the question to avoid technicalities, though I have chosen the simpler course wherever it was found feasible, and have tried to so employ the examples selected that the student who wishes to go further into the matters dealt with may turn to special treatises for further information. For one eminently technical section I ought perhaps to apologise, but the temptation to try and set forth, in concrete form and suitable for the purposes of this book, some account of what is known of the most essential and profound factors concerned in the difficult question of the nature of life and death, health and disease, was great. Probably my apology should go further, and apply to what after all must be failure to explore this mystery to the bottom: my only excuse must be that it may stimulate others to go further.
It was an afterthought to add, in Part I., the considerations on the factors which influence the plant regarded as a living machine, so to speak, in order that the student may the better apprehend the point of view taken of the bearings of the matters discussed in Part II.
With regard to references, it seemed a better plan to give, in the form of notes after each chapter, the titles of the principal books and papers on which a student may base a further course of reading, than to overweight the pages of what is, after all, merely an introductory sketch to a huge subject, with detailed quotations from the numerous sources of information made use of. I have freely expressed my own opinions, but the sources for others are, I hope, as freely given. It will, however, be understood that I have not aimed at a complete bibliography, and, particularly, I have only given foreign references where it seemed that adequate treatment of the subject could not be found in English.
My sincere thanks are due to Mr. F. Darwin, F.R.S., who has kindly looked through many of the proofs, and given me the benefit of several suggestions: and to my wife for the very material aid she has afforded me in the preparation of the index.
H. MARSHALL WARD.
Cambridge,
November, 1900.
PART I.
SOME FACTORS
CHAPTER I.
THE PLANT AND ITS SURROUNDINGS
The plant the central object of study—soil, climate, atmosphere, etc., are factors of its environment. Agricultural chemistry. The plant a machine. Physiology.
If I were asked to sum up the most important result of the numerous advances made during the past decade in agriculture and forestry, I should reply—the clearer and wider recognition of the fact that the plant itself is the centre of the subject, and not the soil, climate, season, or other factors of its environment. Until comparatively recent times it was the habit of farmers, foresters, planters, and gardeners, all the world over, to look upon the plant as a mere item or as a mysterious if important one in their calculations, and to regard the soil as the chief factor in their studies.
Now all is changing, and the world is gradually awakening more and more to the recognition of the truth that the soil and the clouds and the atmosphere are merely reservoirs of more or less inert materials, from which the living plant draws its supplies, and works them up, by means of energy focussed from the sun, into new plant substance.
In other words, the more far-seeing pioneers of scientific agriculture and forestry, etc., are recognising that agricultural chemistry is not the be-all and end-all of agricultural science; but that, in place of the study of the chemical analyses of dead soil, water, air, and plant-remains, which has so long held sway, largely owing, I think, to the influence of Liebig, the student should have his attention more concentrated on the living plant itself and on the physiological actions which make up its life. He must regard the living plant as a sort of working machine—infinitely more complex than any machine made by man, but a machine nevertheless—the purpose of which is to store up energy from the sun, and so to add to our wealth on this planet, at the expense of the extra-terrestrial universe.
It is not, be it noted, that the new study proposes to ignore or abandon the old studies: modern physiology owes too much to the physics and chemistry on which it is partly based, and to the labours of De Saussure, Ingenhousz, Priestley, and others, for that. But it is that the new study recognises that the central point, to which all views must be focussed, is not the one that it was formerly supposed to be. The student is still taught that the chemistry of soils yields valuable information, and that lessons of importance are derived from comparisons of the analyses of the ashes, etc., of plants; but he is no longer able to cherish the hope, however remotely, that such studies solve his most important problems.
The scene—or rather the point to which attention is now directed—is the living, working, energy-accumulating plant itself, and not the dead store of materials in the soil. The reason for the change is not far to seek: it is due to the enormous strides made in the study of the physiology of plants during the last quarter of a century, and the subject abounds in examples illustrating the marvellous advances that have been made, and at the same time showing how, in the progress of researches, made for their own sake—i.e. in pursuit of satisfaction for the intense curiosity of the scientific man—all kinds of side issues turn up which prove to be of value in practice, and suggestive of further thinking.
At the beginning of the nineteenth century—i.e. about 1820—the best thinkers were giving up the old ideas that the environment supplied food, as such, to plants, and had recognised that the plant takes up substances from without and rearranges these in its own body.
The next twenty years or so form a very dark interval in plant physiology, chiefly owing to the influence of the assumption of a special "vital force," an assumption which was not allowed merely to serve as a hypothesis put forward to stimulate research and suggest better ideas, but which gained a hold over men's powers of reasoning to an extent which now appears monstrous and phenomenal.
Many errors crept in during this reign of terror, one of the most fatal of which was De Candolle's revival of the idea of "spongioles"; and another, equally disastrous in many of its effects, was the conception of a sort of vegetable food-extract, humus, existing in the soil in a form peculiarly suitable for direct use by plants. It was during this period that the confusion between the processes of respiration and carbon-dioxide assimilation arose, and exerted its effects for evil into our own day.
The now astounding statement that oxygen-respiration in plants did not occur, laid the foundation of many subsequent difficulties, and so did the positive and authoritative views on the uses of minerals to the plant. Liebig, in fact, stood in the invidious position of being a high authority on purely chemical questions, who was impelled to give opinions on matters which can only be solved by physiological experiments: his great service was to clear up mistakes as regards the chemistry of soils and of plants—his great mistakes were due to his pronouncing on physiological matters; and it may be doubted whether his great services to the purely chemical side of subjects connected with agricultural matters are the more to be admired, or the disastrous influence of his statements on subjects which do not belong to the domain of chemistry should be the more deplored. Be that as it may, he handed on to succeeding generations some weighty errors as regards plant-life, and taught the agriculturist to regard chemical analyses of soils and plant ashes with a reverence which obstructed progress for some time. As a set-off to this we must place his contributions to the destruction of the bugbear vitalism, which was simply preventing enquiry, and his services in bringing together and sifting with power and originality all that had been then acquired as regards the chemistry of the plant, the soil, and the atmosphere.
That Liebig was indispensable in 1840-1850 is one thing; but that his influence should extend to the present day is quite another, and his inevitable mistakes were almost as powerful for future evil, as his clear exposition of the chemistry of his day was productive of immediate good.
Boussingault, working at the same time, 1837-1855, but experimentally with the living plant, taught us more about these matters than any investigator of the time, though it is very probable that the stimulus of Liebig's speculations, good and bad, had its effect in impelling Boussingault to devote his splendid methods to problems of plant-nutrition. Boussingault's contributions to our knowledge of the composition of the dead plant cannot be over-estimated; but he did more than this, because he so clearly apprehended the necessity for asking his questions directly of the living plant, instead of deducing from chemical principles what might be supposed to occur in it; and although future researches showed that even so careful an investigator solved a problem of first importance—viz. the question of the fixation of free nitrogen—the wrong way, it will be found that so far as he did go his conclusions were sound, and well calculated to inspire the confidence with which the world received them. As we are here concerned more especially with the botany of agriculture, however, it is unnecessary to dwell longer on these matters, or on the similar and even more extensive experiments, of world-wide reputation, carried on for so many years, and still being carried on under the liberal auspices of Sir John Lawes, at Rothamsted. Moreover it may be necessary to return to some of these points later on.
Notes to Chapter I
The reader will find a further general account of these matters in Sachs' Lectures on the Physiology of Plants, especially Lectures I. and XII., Engl. ed., Oxford, 1887. He may then proceed to Pfeffer's Physiology of Plants, Engl. ed., 1899, chapter I., and to the account of the history of the subject in Sachs' History of Botany, Oxford, 1890, especially pp. 359-375 and 445-524. References to more special literature will be found in Pfeffer.
CHAPTER II.
THE PLANT AND ITS FOOD
The food of plants—"Vital force"—Other errors—Liebig and Boussingault—The botany of agriculture. The synthesis of carbohydrates—The physiology of plant-nutrition. The persistence of misconceptions.
The year 1860 may be regarded as a landmark of importance in the history of plant physiology, for it was in that year that Sachs discovered that the bringing together of water and carbon-dioxide, in the green chlorophyll-corpuscles of the plant exposed to sunlight, results in the formation of the grains of starch found in these corpuscles.
Previous to this date Dutrochet (1826-37) had introduced the then crude idea of osmosis into physiology; vegetable anatomy had improved, and the modern conceptions of the living cell, protoplasm, nucleus, etc., were slowly looming; sieve-tubes had been discovered, and the proteids and starch in various parts of the plant examined; and the suggestion was abroad, replacing Liebig's idea that plant acids were the first products of carbon-assimilation, that some substance, of a slimy nature, was manufactured in the cells of the leaves and thence distributed as the formative material from which the plant constructed its parts. Davy and Boussingault had even surmised that a carbohydrate might be the first-formed product in assimilation.
There can be little doubt that Sachs' classical proof, by direct physiological observation and experiment, first brought forward the truth of organic synthesis in the plant in a concrete and convincing form.
But it did more than that. It laid the foundation of the modern physiology of plant-nutrition on ground already prepared by De Saussure and the earlier workers; for, in addition to emphasising the truth of organic synthesis—a truth which had been gradually impressing itself on the world for some years—Sachs' discovery showed clearly the real meaning of carbon-assimilation as a process for obtaining combustible food, which the plant then proceeds to make use of.
Many points were rapidly cleared up at once, or if not explained were at least put into a strong light for further enquiry, and plant-nutrition soon ceased to be the mysterious subject for all kinds of wild conjectures that it had hitherto been.
The meaning of thin leaves, with numerous stomata and finely ramified or divided vascular bundles, became more apparent, as also did the significance of the ascending transpiration current; the storage of starch-grains in tubers, medullary rays, roots, seeds, etc., obtained meanings not understood before; the spread of roots in the soil, and the gradually discovered properties of the finer rootlets and of the root-hairs, fitted naturally into their places; and, in short, a thousand facts, otherwise isolated, became collated into an intelligible system, full of suggestions for new work, such as has since gone on and is now being pursued with an activity and success never before realised in the history of science.
As time went on, while the general truth of Sachs' views was confirmed, a number of detailed discoveries were made which seemed to contradict them in certain points. It was found that not all leaves form starch, for some contain sugar or oil; but Holle and Godlewski proved experimentally that this oil may be replaced by starch if the conditions of assimilation are slightly modified. More recently Hébert discovered that the stalks and leaves of grasses contain a peculiar form of gum, which was formerly confounded with starch, a substance not abundant in them. Then came Schimper's discovery of starch-forming corpuscles, which, if supplied with sugar, are able to form starch-grains in the dark, as in tubers, etc., underground; and as subsequent researches have proved that the chlorophyll-corpuscles—which are morphologically the same as the starch-forming corpuscles and can be replaced by them—are also able to form starch-grains from sugar, as proved by the experiments of Boehm, Acton, Meyer, Laurent, Bokorny, Saposchnikoff, and others, it soon became evident that nothing essential needed altering in Sachs' view that starch is the first visible product of carbon-dioxide assimilation, only it became clearer that the starch-grains are built up by the protoplasm from glucose or some similar body, and represent so many packets of reserve materials put by for the present because not required for the immediate needs of the cell.
Boussingault showed, about thirty years ago, that assimilation soon stops in green leaves if cut off from the plant, not because the leaves die, but owing to some "maximum capacity" being attained. Sachs had shown that the starch passes down to other parts of the plant in solution as glucose.
Neither time nor space will permit me to go into the enormous field of research and results opened up by these and similar observations made between 1860-70. It must suffice to say that they led to the discovery and study of the diastatic and other enzymes in the leaves and other green parts of plants, and to a clearer understanding of what was already known of them in seeds, and this knowledge reacted at once on our insight into the processes of transport of reserve materials and constructive materials from one part of the plant to another, matters which will be referred to later on.
It remains to explain Boussingault's difficulty as regards the cessation of assimilation. Recent researches confirm the view that at least three causes are at work to bring about the inhibition of the carbon-assimilation: first, the chlorophyll-corpuscles become filled to excess with starch, which cannot get away because all the passages are full and the products are inhibiting the further action of the enzymes which should dissolve the solid granules; secondly, the leaf being detached from the plant explains why the soluble products cannot get away, for this makes a great difference in the rate of exhaustion of the leaf; and, thirdly, the same fact involves that the leaf can obtain no further supply of salts of potassium, etc., without which elements the processes in question cannot go on.
These and numerous other deeper insights into the process of assimilation, obviously strengthen the force of Sachs' discovery; though it by no means necessarily follows that starch-grains are always the resting form of the products of assimilation, and we now know that such is often not the case: we now have much deeper glimpses into the initial products of carbon-assimilation than Sachs had in 1860, but this enhances rather than detracts from the importance of his splendidly worked-out discovery. Put more generally, we may now say that the process of carbon-dioxide assimilation in green leaves under the influence of light is a process of synthesis—photo-synthesis—resulting in the building up of a carbohydrate such as sugar, inulin or starch from the elements carbon, hydrogen and oxygen.
But it must not be supposed that the importance of Sachs' discovery, and the rapid consequent extensions of our knowledge, did their work forthwith in disabusing men's minds of old and erroneous notions. To say nothing of numerous smaller misconceptions which still held their ground owing to the stupendous ignorance of plant-physiology which prevailed, we find incompetent teachers and text-books were still propagating ideas worthy of ancient times. The confusion between oxygen-respiration and the gas interchanges in carbon-assimilation was by no means eliminated even recently, though it can no longer withstand the deliberate onslaughts now made on it. That the roots take up food as such from the soil, and that that food is directly employed by the plant for its nutrition is even yet implied in daily conversation around us; and although matters have advanced so far that everyone now knows that the substances at the roots must be in solution, ere they can be received into the plant, it sometimes leads to astonishing replies, if we press the question very far as to how the absorption takes place, in an elementary examination of agricultural students. That manures are foods to the plant, that sap circulates, that transpiration is of use to keep the plant cool, and wood is a "porous body," etc., are only a few of the misconceptions still current, in a decade that has found publishers for a work advocating that roots are congealed sap, and that the leaves of plants absorb the moisture and dust of the air, and so provide the plant with food, and for a paper explaining the action of root-hairs as tubes with open pores at their tips. But the gravest misapprehensions current among us are due to the crude ideas as to what a plant really is: this, I take it, is owing to the difficulty of grasping what physiologists mean by organised structure, and leads to regarding the living being either as a mere aggregation of chemical compounds, built up by the ordinary play of chemical forces, as we know them, acting on dead matter, or, as in the days before organic chemistry, as a mysterious entity endowed with "vital force," and with properties not amenable to scientific investigation. The mistaken notions as to the powers of roots to "select" those substances which the plant requires, and to reject useless ones was merely an expression of this belief.
The rock on which all are liable to come to grief—the chemist or physicist who requires all his facts in terms of analyses and proportions by weight, and therefore takes too mechanical a view of the subject, or the man who is not scientifically trained at all, and therefore is more liable to go to the other extreme and regard the plant as a mysterious something which grows and has poetical associations and traditions—is the great fact of organised structure, and it is the recognition of this fact and some of its consequences which has altered the whole position of the subject, and brought the study of the plant into the domain of physiology. The living plant, its structure and organisation, the functions of its mechanism, and its relations to the environment, thus forms a subject apart from that which concerns the chemical composition of the plant and its environment, and this distinction designates, in a word, as it were, the change which has been brought about by modern biology.
A point to be emphasised to the utmost where agricultural students are concerned is that the essential process of feeding is the same in a green plant, a fungus, and an animal; the greatest confusion still exists with regard to this matter, owing to misconceptions as to the real meaning of the functions of the chlorophyll-corpuscles when supplied with carbon-dioxide and water and the energy of the sun's rays. The plant does not feed on carbon-dioxide, any more than it feeds on oxygen—it feeds on the organic material after it has been constructed, and the chlorophyll-function is merely one mode of obtaining supplies of such organic substance.
