Biolopcnl ,Journal of the Linnean Society (1987), 32: 5-16 What is Ecology? Professor Sir A R T H U R TANSLEY, F.R.S. Chairman of the Xature Conservany and President of the Council for the Promotion of Field Sludies (Originally published in 1951 as a pamphlet by the Council for the Promotion of Field Studies, which became the Field Studies Council) The word ecology is now appearing in print more and more frequently, even in some newspapers. But how many people have any clear idea of what it means? Apart from those who actually work a t the subject, very few indeed! No doubt the great majority of biologists, with no special interest in ecology, have a general notion of what it is about, but many of them are far from appreciating the full scope and potentialities of the subject. One is often asked just what ecology is, and an attempt to make its nature and aims as clear as possible to all who are or may be interested is well worth while, for active ecological work of one kind and another will be of vital importance to the whole human race in coming decades. EARLY HISTORY The word, or rather its German equivalent, was actually invented by Ernst Haeckel as long ago as 1870. Haeckel was a well-known German professor of zoology who gained fame, and indeed notoriety, by expounding and defending the theory of evolution, which was a t that time far from being as widely accepted as it is to-day, and joining with it a philosophy of atheistic monism. All that does not concern us at all. The word was very little used during the next 25 years, and it was not till 1895 that Eugenius Warming, Professor of Botany at Copenhagen, published a book in which the Danish equivalent of the adjective “ ecological” was used in the title. This may be rendered into English as PlantAggregates: Foundations o f Ecological Plant Geography. In the following year a German translation was published, and this was widely read in England and America, most English-speaking professional botanists being able to read German hut not Danish. Warming’s book was an excellent and very attractive exposition of one fundamental aspect of its subject. It opened something like a new world to many young botanists, and its publication marks the beginning of the general use of the words “ecology” and “ecological.” Shortly afterwards A. F. W. Schimper in Germany, and Charles Flahault in France, who had long occupied themselves with these topics, also wrote on the subject from somewhat different angles, 5 0 1951 Field Studies Council 6 A. TANSLEY Schimper dealing with what was then known of the vegetation of the whole world in a large and very fully illustrated volume, of which an English translation appeared a few years later. The publication of Warming’s and Schimper’s books was a tremendous stimulus to ecological work in America and England. A good many of the papers published during the first years of the new century were, however, rather trivial and some of them decidedly slovenly. Besides stimulating many good biological minds, ecology had a great attraction for weaker students, because it was so easy to describe particular bits of vegetation in a superficial way, tending to bring the subject into disrepute. Notable exceptions to this futility were the works of the great pioneers of scientific plant ecology in America, H. C. Cowles and F. E. Clements, and their pupils and followers. I n Britain the pioneer work was done by Robert and William G. Smith (Robert Smith had been a pupil of Flahault a t Montpellier) and their pupils and co-workers, first in Scotland and then in England. In 1904 a small body of enthusiastic workers in plant ecology formed the “British Vegetation Committee,” succeeded in 1913 by the British Ecological Society with its Journal o f Ecology; the Journal o f Animal Ecology was started by the Society in 1932. In 1915 the Ecological Society of America was founded, and its journal, Ecology, began to appear in 1920. The reader may well complain at this point that he has still not been told what ecology actually is. The excuse must be that some little historical background was desirable first. It will already have become apparent that plant ecology has to do with vegetation and with plant geography. But there is much more to ecology than that. THE ESSENCE OF ECOLOGY The word ecology is derived, like the common English word economy, from the Greek oikos, a house, abode, dwelling, and this etymology gives the key to the meaning of the word. I n its most general sense ecology may be defined as the study of plants and animals as they live in their natural homes, and this implies the study of all they are and do when they are “at home.” I n the case of mobile animals “home” must be extended to include all the places they are in when they are feeding, seeking food, escaping, sheltering, migrating or breeding-in fact the whole of their behaviour during life. Most plants do not move actively about, but they also “behave,” in the sense that they grow in various directions, and, to various extents, make their own food, and reproduce themselves. While modern ecological work at the end of the last century and the beginning of this was practically confined to plants, it was soon extended to animals, and animal ecology is now an actively developing and important subject. In fact anything like a satisfactory study of the ecology of the populations of living beings inhabiting a particular area must include both plants and animals because they are not only constantly affecting one another, but often depend upon one another for their wellbeing or even their continued existence. Green plants form the basis of life as it is lived upon the earth because they, almost alone, are able to make organic matter, i.e., the substances of which all living organisms consist, out of inorganic materials like water, carbon dioxide (from the air) and mineral salts (from the soil). Animals depend for their food on this activity of green plants, directly through eating them (herbivorous WHAT IS ECOLOGY? 7 animals), or (carnivorous animals) indirectly by eating other animals which have fed on the plants. THE PLANT CARPET And there is another sense in which plants form the basis of life upon the earth. I n favourable conditions of climate and soil plants form a more or less continuous carpet on the earth’s surface-woodland, grassland, heath, moor, bog, etc.-where the carpet has not been destroyed by man or by some catastrophic natural agency. I n very unfavourable climates, where there is insufficient water, intense cold throughout the year, or constant violent winds, plants are unable to form a continuous carpet. In subtropical (and other) deserts the kinds of plants that can live there at all are represented by scattered individuals with bare ground between them. In the severest conditions of all (e.g., on icefields or in the immediate neighbourhood of volcanoes) plants may be absent altogether. The normal plant carpet not only supplies the food of herbivorous animals on which the carnivores depend for their food, it also provides the shelter which most animals need because many of them cannot live, at any rate continuously, completely exposed to the elements. Others require shelter to hide from their enemies, or again to conceal themselves from intended prey so as not to frighten the intended victim away before the pounce can be made. There are thousands of kinds of small “invertebrate” animals-many insects and tiny creatures belonging to other groups-which live in the carpet of vegetation or in the soil on which it grows, and never leave it. Plants thus perform essential services to animals by supplying them with food and shelter; herbivorous animals destroy plants by eating them, but some (mainly insects) also benefit plants by bringing pollen (essential for reproduction) from flower to flower, and many animals of all kinds and sizes also help to distribute seeds from place to place. PLANT COMMUNITIES The plant carpet differs, of course, very much according to the conditions in which it grows, in the first place climate and to a large extent also soil. We have only to think of the luxuriant evergreen tropical rain forests such as we see in Malaya and neighbouring regions, the deciduous temperate forests of western and central Europe and eastern North America, the leathery small-leaved scrub of the Mediterranean region, and the northern and sub-alpine needle-leaved coniferous forest, to realise the primary importance of climate in determining the type of forest that can exist in a given region of the world. And to the forests we must add the great natural grasslands-the American prairies, the south Russian steppes, the South African veld-determined by yet another type of climate, with summer rainfall but drier than the forest climates. T h e subtropical deserts of North Africa and south-west Asia are much drier still, with constant deficiency of water and very sparse distinctive vegetation. Then there is the special vegetation of high mountains. The influence of soil conditions is seen in the characteristic vegetation of constantly water-logged soils, giving us swamps, marshes, fens and bogs, each with different kinds of plants, and at the other extreme the vegetation of very dry sandy soils. I n forest regions, too, different kinds of tree often live on different types of soil. 8 A. ’I‘ANSLEY The plant carpet of the world falls naturally, in fact, into unils of vegetation, or, as we call them, plant communities, each with its own kinds or species of plants, of which the dominant form and structure are distinctive of the community, and have a dose relation to the climate or soil, or both, in which they occur. It was these units of vegetation or communities of plants that Warming and Schimper described and analysed in their epoch-making books. (Many of the great types of vegetation had, of course, been described much earlier, e.g., by the great geographer and naturalist, Humboldt.) INTEGRATION AND ORGANISATION OF COMMUNITIES M‘e speak of plant communities by analogy with human communities, for when any set of animals or plants live side by side in a particular habitat they always tend to form an entity, primarily because they are living under more or less identical conditions (differing from those of adjacent areas) which have similar (though not necessarily identical) effects upon them all, and also because the individual plants affect one another, through competition for living space and often in other ways too, as we shall see presently. Of course, the bonds uniting the members of a human community are very much closer than those of a plant community or of most animal communities- the “integration” is much greater-because human beings can communicate with one another through speech and thus are in a position to have profound psychological effects on one another’s lives-apart from the unpleasant habit of imprisoning, torturing or killing one another in large numbers. T h e interactions of human beings living together have led to a high degree of organisation in human communities, and this has become extremely complex in those which we call civilised. Considerable organisation is found also in the communities of many gregarious non-human animals, for example in some herbivores (cattle, deer, antelopes) and such carnivores as wolves. But it is in certain gregarious insects (e.g., hivebees, termites and ants) that we meet with the most complex community organisation in non-human animals, with far-reaching structural and functional differentiation and division of labour between different “castes)’ (workers, “soldiers,” etc., which are structurally incapable of breeding, besides the comparatively few sexual individuals) , and a completeness of “integration” of the whole community which is still (fortunately) not found among human beings, though it seems to be the ideal of some moderns, and certain approaches to it have been made from time to time in human history. From their very nature plants cannot bring about the kinds of organisation and integration of the community which have just been mentioned. The members of human and of many animal communities typically belong to a single kind or species of animal, though there are a certain number in which two or more species are associated. But plant communities typically consist of several or many different species living together-in the most complex, such as the Malayan rain-forest, several hundred species in one community. Between the different species of a plant community various relations exist. Some compete directly with one another for living space, water and light, and the wcakcr individuals go to the wall. But in complex communities such as forests in which the vegetation is “stratified,” there is no direct competition between the different layers or strata of the forest (trees, shrubs and herbs). T h e shrubs do WHAT IS ECOLOGY? 9 not compete with the trees, nor the herbs of the forest floor with either trees or shrubs. O n the contrary, many of the herbs of the forest floor could not exist except in the shade provided by the foliage of the tree canopy which makes the air relatively quiet and damp inside the forest, and some of them depend on the humus of the soil formed by the fallen and decaying leaves of the trees. Then, again, the great trees of a tropical rain forest support a number of woody climbers (lianes) which climb up the trunks and flower on the forest roof, and also a number of plants (epiphytes) which are not rooted in the soil a t all but grow upon the trunks and branches of the trees and draw their food from humus lodged in crevices of the tree bark and their water partly from this and partly from the abundant rainwater running down the trunks of the trees. Quite similar relationships, though on a much less elaborate scale, exist between the different components of temperate forests. All such relations of dependence of some members of a plant community upon others build up the unity of the community as a whole-a unity which is originally based on the fact that it consists of a particular collection of species growing together in the same general conditions of climate and soil, but is notably integrated by the establishment of any form of interdependence between the members. Practically all plant communities have animals associated with them in different ways, ranging from species of birds and mammals which live in forest, or on grassland or heath, to minute invertebrates which are closely tied to the plants (often to particular species of plant) by the exigencies of their lives. Sometimes there are small local communities of small vertebrates, such as rabbits or voles, in our own woodlands or grasslands, or of various invertebrates, within the general plant community; and these may be very highly integrated in themselves-ants’ nests in a conifer (or other) wood are a good example. Many species of the larger animals, mostly birds and mammals, which are highly mobile, commonly range, however, through many plant communities, and are not tied to, or in any way dependent upon particular ones. But when you have scattered individual animals, or communities of animals, which are so tied, the entire complex of natural plant and animal life (so long as it is not interfered with by man), together with the physical factors of climate and soil which permit its existence, forms, when mature, a n integrated and balanced “system” which may be called an ecosystem, and can sometimes maintain itself, apparently indefinitely, so long as the conditions which determine it continue. Such is the subject matter of ecology and it will be realised at once that it is both extremely wide and extremely complex. COLLABORAIION IN ECOLOGICAL WORK To get really satisfactory ecological knowledge of an ecosystem it is necessary to make a thorough study of both the plants and the animals present and to determine their relations to one another, as well as the nature of the habitat or “home” of the community and the relations of the organisms to it. Unfortunately, in current biological training the student specialises either in botany or in zoology and has rarely sufficient knowledge of the other subject to enable him to make a comprehensive study of both plants and animals. I t is true that animal ecologists are more and more being forced to acquire some considerable acquaintance with the plant communities on which their animals 10 A. TANSLEY depend, b u t it is still rare for a plant ecologist to have enough zoological knowledge to make an adequate ecological investigation of the animal elements in a n ecosystem. Close collaboration between an animal and a plant ecologist, or by a team of several, is one way out of this difficulty. But such collaboration, to be really successful, must be genuinelyjoint work in which each partner fully follows and understands the contributions of the others. Only thus can a unijied picture of the whole be built up. It was the fresh emphasis on the natural aggregates that plants form when left to themselves, and on the dependence of these on the conditions under which they live, that led to the great modern development of ecology which began about the turn of the century. Of course the great types of vegetation had been recognised and given names in common language from the earliest times. It could not have been otherwise, since they were the most conspicuous and important parts of primitive man’s environment even after he had begun to cultivate the soil and to keep his flocks and herds. They continued to be so through the ages till the enormous growth of great cities eventually separated the people who lived in them from their original surroundings-a fateful divorce whose price we are now paying. But recognition and even intimate practical familiarity with the great types of plant community and the animals they contain is a very different thing from their use as objects of close scientific study; and it is the increasing appreciation of their profound interest and value from the scientific and the practical points of view that has marked the past halfcentury. AUTECOLOGY AND ANCILLARY STUDIES Close study of a plant community always leads to questions about the r61e and behaviour of particular species which are members of it. Investigation of the ecology of individual species (autecology as it is called), especially of the dominant (usually, but not always, the tallest) members of a community, is an essential (and the most time-consuming) part of ecological work. Until the most important facts about them are learned it is impossible to penetrate at all deeply into the ecology of the community as a whole, because of course the latter is made up of the behaviour and interactions of the species comprising it, and the dominant species (e.g., the trees that form the canopy of a forest, the ling or heather of a British heath, certain kinds of grass in a meadow, etc.) may have a controlling influence on the whole community. While the great development of modern ecology was initiated, as we have seen, by the recognition of the importance of plant communities, and they have remained a leading subject in current research, the ecology of single species presents the most fundamental problems of all. This kind of fundamental ecology involves a knowledge and use of the most various branches of biology. First of all, we must know our plants and animals, recognise the named species and varieties to which they belong, and that is “systematic” botany and zoology, or taxonomy. Then we must understand the members or organs of their bodies and how they develop in their life histories, which is morphology. We cannot understand how a plant or animal works in detail unless we are familiar with the structure of the tissues (cellaggregations) of which it is composed and through which its activities are carried on (histoloo). T h e study of how an organism does work, both its internal WHAT IS ECOLOGY? 12 processes and how it acts on its environment, and its environment on it, is physiology, a science which underlies most ecological phenomena. Again, we must know something of the modern science of heredity, called genetics, for many of the forms we meet in the field cannot be understood without knowing or finding out their genetic status within a species, with which their ability to live in a particular community may vary. It is not intended to suggest that every good ecologist must be an expert in all these branches of biology-nowadays an impossible demand. Some sound elementary knowledge of all of them he must have, but that is not so difficult to acquire. Above all, however, he must have practical experience of field workand the more the better. Ecology is essentially a “field subject”: progress cannot be made in it without constant observation (and frequently experiment) in the field, though some of the ecological problems met with in the field have to be attacked in the experimental garden or the laboratory, or both. Besides the particular branches of biology referred to there are certain nonbiological subjects of great importance to the ecologist-climatology and meteorology, geology, physiography and soil science (pedology). This is because these subjects deal with climate and soil, which, as we saw, are the main primary factors determining the distribution of plant communities, and training in them is very useful to the ecologist. I n default of that he must make himself acquainted with those parts of them which are clearly important in the particular work on which he is engaged. All this may sound very alarming to the beginner who is attracted to ecological work but is doubtful of his capacity to fit himself for it. There is, however, a consideration which should be reassuring. While an ideally complete equipment for ecological work is impossible in the time available to the student, and indeed beyond the mental capacity of any individual, the field open to research is so vast and the problems it presents so diverse and of such varying degrees of difficulty that everyone who wants to work in the field can find something really useful to do, whatever his natural bent and his training. And he can pick up a good deal of necessary knowledge as he goes along. ECOLOGY AND NATURAL HISTORY In the early years of the century ecology had a hard fight for recognition, at least in this country-it was more quickly and widely accepted in America. A good many biologists tended to regard the subject with something like contempt-an attitude partly excused by some of the superficial, trivial, or slovenly observations published under its name. “There is nothing new in all this,” they would say, “it is only the old natural history masquerading under a high-sounding name-and not always very good natural history at that!” It has needed some decades of steady work to change this attitude, and even to-day a friend of mine, a first-rate biologist in his own line, professes not to know what ecology means. But there is plenty of evidence now of its wide acceptance in the schools and universities and of the increased recognition of its scientific and practical importance. It is perfectly true that ecology is natural history-it has been aptly called scientific or systematised natural history-and that a great deal of the excellent 12 A. TANSLEY natural history work done before Warming and Schimper would now be classed as ecology. The growing popularity of the subject in this century represents a breakaway from the confinement of serious work in biology to the laboratory, museum and herbarium, which at one time divorced the study of plants and animals from their actual life in their natural surroundings. Laboratory, muscum and herbarium are still, of course, and always will be, absolutely necessary for the pursuit of biology, not least as adjuncts to the study of ec,ology itself, but the primary place in which to work at ecology is necessarily the field. APPLIED ECOLOGY What has been written up to this point relates to the pure science of ecology, pursued for the attainment of fundamental knowledge about living nature and her ways. But, as in most sciences, we have to regard also the applications of’ such knowledge to practical human uses. Just as a fundamental knowledge of physics and chemistry is necessary for engineering and for modern industrial processes, and of human anatomy and physiology for surgery and medicine, so a knowledge of outdoor biology, soil science and climatology, which may be best integrated and expressed as ecology, is necessary for any industry concerned with growing plants, such as agriculture and forestry--for all use of land which contemplates maintenance of the existing natural vegetation or its replacement by some crop of man’s choosing. For these purposes it is not always necessary to take account of all the subtleties of pure theoretical ecology, though many of its concepts turn out to be very useful in applied work. A thorough understanding of the ecology of natural forest is a very important preliminary to the work of the forester in dealing with the growth and timber production of artificial plantations of trees, and neglect of this training has led to costly mistakes. The same may be said of the ecology of “rough grazings“ consisting entirely of native plants which were not sown but came there by themselves. In all such cases a thorough knowledge of what nature is doing on a particular site is the best and often essential preliminary to satisfactory human exploitation. An important job in which ecology is highly relevant is the war against p e s k attacking crop plants, of which various fungal and insect parasites are the most important. Here, no doubt, mycology (fungi) and entomology (insects) are the primary subjects necessary, as is now recognised by the appointment of trained mycologists and entomologists as pest-control officers. But mistakes arc frequently made through too narrow a view of the work required. We not only have to understand the structure and life histories of the fungus and insect pests, but also the ecology of the host plants, the different conditions in which they flourish and also those in which they are most liable to attack by the parasite. A comprehensive and penetrating knowledge of the whole background of a pestattack is the soundest basis for devising efficacious measures against it. Again, serious errors in practice may arise from too narrow a concentration o n destruction of a pest attacking a natural community, where the means of’ destruction employed may involve the injury or death of other members of the community that are useful or essential to the wellbeing of the whole, or the destructive agent is found to have other unexpected and undesired results. WHAT IS ECOLOGY? 13 THE GENERAL CASE FOR APPLIED ECOLOGY At the outset it was said that active ecological work of one kind and another will be of vital importance to the whole human race in coming decades, and the reader will naturally want to know what are the grounds of such a sweeping statement. They may be stated most generally by saying that we cannot control or utilise nature to the best advantage unless we first understand her, and that ecological work is the right road to such an understanding so far as it concerns the vegetation and animal populations of the world. Apart from the most sterile areas and those occupied by human constructions the land surface of the earth is covered either by natural vegetation-----muchof it largely modified by men’s activities-or by planted crops yielding foodstuffs or industrial raw materialtimber, fibres, rubber and so on. Much natural vegetation is annually destroyed by human agency, intentionally or carelessly, and not replaced, leaving great areas derelict, or the method of cultivation is ill-judged, or an area is abandoned after crop failure. Any of these mistreatments may lead to extensive erosion by rain or wind or to “dust bowl” formation, according to the climate. Such careless and disastrous exploitation of man’s natural heritage has been ably dealt with in more than one recent book. Very little ecology is necessary to point the way to more intelligent treatment of nature in the most flagrant cases of abuse. But there are many others in which a proper understanding of the ecology of the vegetation to be dealt with is really necessary before what is practicable and what is most advantageous can be settled. T h e recent groundnuts fiasco in Tanganyika is a conspicuous case in point. A thorough ecological investigation of the areas, especially of their climate and soil, of the bush already growing there, and of the conditions necessary for the good growth and economical harvesting of ground-nuts themselves should have been carried out before this vast and ill-fated adventure into unknown ecological territory was decided upon. And there were, in fact, highly qualified ecologists available for such a job. Of course, it would have taken time and the promoters of the scheme were in a hurry. But if you try to “rush” nature she is very apt to avenge herself to your ultimate detriment. Nature just won’t be rushed. THE FOOD-POPULAlION PROBLEM In the not very distant future the human race is faced by dangers at least as threatening and certainly more inevitable than those of atom bombs, because they are inherent in the activities of many hundreds of million people, unless necessary counter measures are taken in good time. Attention has been called to them by several instructed and responsible writers, but there is little sign that those responsible for international corporate action have taken heed. The human race, or rather, some of the largest parts of it, is breeding a t a rate that will very soon outrun food supplies which are available now or are likely to become available unless very far reaching and drastic steps are taken at once or in the near future. The possible steps seem to be three-fold. The first is evidently reduction of the birthrate of the more prolific races. This is clearly a problem of very great difficulty but one that ought to be squarely faced, for example by the Indian Government. It scarcely concerns the subject of this pamphlet though it is 14 A. TANSLEY doubtless part of what is now often called “human ecology,” which is far too vast a subject to deal with here. This is certainly the most important of all counter measures, for unless the present rate of population growth is brought down to a much lower level it would seem that no increased annual supply of food which will be at all practicable could possibly feed the future population of the world. Vital as reduction of birthrate is, it is also most important to increase the quantity and improve the quality of the world’s food supply, in the first place by getting greater and better quality yields from food-producing land that already exists. This is essentially a job for the scientific agriculturist; scientific agriculture is certainly applied ecology, but it is clearly ecology of a very special kind, depending on a profound knowledge of the optimum conditions for cultivation of a particular crop in the particular district where it is to be grown. Quantity of yield depends partly, and quality mainly, on breeding the best new strains of food plant and that is work for the geneticist or plant breeder. The new wheats bred in England 40 years or so ago are only one of the earliest and most conspicuous examples of the striking successes which have been reached in this field. There have been many others since. A third step is to bring more potentially food-producing land under cultivation. That this could be done on a large scale in some tropical and subtropical countries of at least three continents is certain, but in view of the urgent need of much more and much better food it is necessary for the proposed sites and proposed crops to be chosen only with the fullest possible knowledge that can be obtained beforehand and tested by carefully planned field experiments. The ecology of existing vegetation on possible sites has first to be thoroughly studied and compared with that of possible food plants, and then a series of crucial experiments undertaken before any schemes on a large scale are entered upon. I n this way very considerable successes could no doubt be obtained and the world’s food supply significantly increased. This is essentially a job for ecologists and ecologically trained agriculturists, and it is one of the most important jobs in the world, Many ancillary tasks would have to be undertaken, of course: improved methods of cultivation, of harvesting, of transport, marketing and distribution, and training of natives in suitably adapted modern agricultural practice. If it is to be done on a sufficient scale the work must probably be to a large extent mechanised. Hurry is fatal, as we have recently seen, but delay will be equally fatal, for the danger of widespread starvation is creeping upon the world year by year. It is not far from India to-day; serious malnutrition and under-nutrition, already rife there, is increasing, and the importation of food from other countries is a mere temporary palliative. The existing food resources of the world are simply not large enough to meet the need. An ecological approach to these problems is a necessity because they all ultimately depend on ecology, whatever specialised technical training may be required in addition. The means of acquiring the ecological outlook are therefore of prime importance. ’1 WO RECENT ORGANISATIONS Within the last few years two organisations, both of which have begun to serve the ends of ecology, have been set up in this country. One is the Nature WHAT IS ECOLOGY? 15 Conservancy, which is a minor Government Department under the same committee of the Privy Council as the Agricultural Research Council. The Conservancy itself, which makes its own policy and conducts its own affairs, reporting to the Lord President, who is responsible for it to Parliament, every year, is a body of about 20 people, mainly naturalists. The work of the salaried scientific staff is very largely ecological. I n the National Nature Reserves set up by the Conservancy, good samples of the different natural plant communities are to be preserved, together with typical animal populations, and in the Reserves ecological surveys and research will be carried out by members of the staff and other qualified workers. I n this way a body of ecological knowledge will be amassed which will be at the disposal of Government Departments, public authorities, and all responsible bodies and individuals who can use it. Meanwhile British ecology should be notably advanced. While considerable knowledge of it has been built up during the past half-century there is a great deal more that we do not yet know about our native vegetation and animals. All such knowledge will be valuable not only as pure science but often as a guide to the wisest forms of land utilisation. The other organisation is the Council for the Promotion of Field Studies, which is a private body, but of national scope. This has established four residential “Field Centres” each under the charge of a trained warden and assistant warden. T o these Centres students come, classes from the upper forms of schools with their own teachers, university students, and private individual students, for a week or more at a time, to carry out field work of every sortmuch of it ecological-under the expert guidance and sometimes the personal tuition of the warden or assistant warden. The four Centres are widely dispersed in England and Wales: all four are situated in unspoiled rural country of very different types, so that a considerable variety of plant communities and animal populations can be studied by visiting two or more in succession. The work of the Council has been very successful and has aroused a great deal of enthusiasm among the visiting students. Not only does it provide training in an attractive subject of vital importance for the future, but by taking the students into the open air and into unspoiled nature as the primary field of work it has a unique effect on their minds and bodies. For many it is their first experience of a new and fascinating realm of mental activity-both scientific and delightful. While the objects of the Council are primarily educational, many schemes of ecological research are in progress or projected at the different Centres, and this is a vital element in their work. Unfortunately the Council’s work is still hampered by insufficient means. The cost of the scientific staffs at the Centres cannot be met from students’ fees, and Government grants that have enabled the Council to carry on up till now are to be drastically reduced. CONCLUSION Those who have had no actual experience of serious ecological work in the field, and who want to understand its essential nature, must realise that this lies in a distinctive point o f view in the approach t o j e l d biology. The ecological approach comprehends the diverse features of the landscape with their progressive change or relative stability, the factors of climate and soil, whether these are wholly 16 A. TANSLEY natural, as in virgin vegetation, or more or less profoundly modified by past or present human activity-and the living organisms themselves, the centre of the picture, as they are affected by the factors of their environment and by their interactions with one another. When once grasped this characteristic unifying point of view is unmistakable. There are some biologists who are real experts in their own lines of work, but who have never got hold of it at all, though their own specialised knowledge is essential to progress in ecological research. An eminent German plant geographer of the last generation once wrote that the strength of ecology depends upon the enormous wealth of data that pour into it from every side, and it was this specialised knowledge from the different branches of biology and from the ancillary sciences that he had i n mind. But it is the unifying point of view that makes ecology what it is.
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