Ancient Plants – Conclusion

CHAPTER XIX
CONCLUSION

In the stupendous pageant of living things which moves through creation, the plants have a place unique and vitally important. Yet so quietly and so slowly do they live and move that we in our hasty motion often forget that they, equally with ourselves, belong to the living and evolving organisms. When we look at the relative structures of plants divided by long intervals of time we can recognize the progress they make; and this is what we do in the study of fossil botany. We can place the salient features of the flora of Palæozoic and Mesozoic eras in a few pages of print, and the contrast becomes surprising. But the actual distance in time between these two types of plants is immense, and must have extended over several million years; indeed to speak of years becomes meaningless, for the duration of the periods must have been so vast that they pass beyond our mental grasp. In these periods we find a contrast in the characters of the plants as striking as that in the characters of the animals. Whole families died out, and new ones arose of more complex and advanced organization. But in height and girth there is little difference between the earliest and the latest trees; there seems a limit to the possible size of plants on this planet, as there is to that of animals, the height of mountains, or the depth of the sea. The “higher plants” are often less massive and less in height than the lower:Man is less in stature than was the Dinosaur:and though by no legitimate stretch of the imagination can we speak of brain in plants, there is an unconscious superiority of adaptation by which the more highly organized plants capture the soil they dominate.

It has been noted in the previous chapters that so far back as the Coal Measure period the vegetative parts of plants were in many respects similar to those of the present, it was in the reproductive organs that the essential differences lay. Naturally, when a race (as all races do) depends for its very existence on the chain of individuals leading from generation to generation, the most important items in the plant structures must be those mechanisms concerned with reproduction. It is here that we see the most fundamental differences between living and fossil plants, between the higher and the lower of those now living, between the forest trees of the present and the forest trees of the past. The wood of the palæozoic Lycopods was in the quality and extent and origin of its secondary growth comparable with that of higher plants still living to-day:yet in the fruiting organs how vast is the contrast! The Lycopods, with simple cones composed of scales in whose huge sporangia were simple single-celled spores; the flowering plants, with male and female sharply contrasted yet growing in the same cone (one can legitimately compare a flower with a cone), surrounded by specially coloured and protective scales, and with the “spore” in the tissue of the young seed so modified and changed that it is only in a technical sense that comparison with the Lycopod spore is possible.

To study the minute details of fossil plants it is necessary to have an elaborate training in the structure of living ones. In the preceding chapters only the salient features have been considered, so that from them we can only glean a knowledge similar to the picture of a house by a Japanese artist:a thing of few lines.

Even from the facts brought together in these short chapters, however, it cannot fail to be evident how large a field fossil botany covers, and with how many subjects it comes in touch. From the minute details of plant anatomy and evolution pure and simple to the climate of departed continents, and from the determination of the geological age of a piece of rock by means of a blackened fern impression on it to the chemical questions of the preservative properties of sea water, all is a part of the study of “fossil botany”.

To bring together the main results of the study in a graphic form is not an easy task, but it is possible to construct a rough diagram giving some indication of the distribution of the chief groups of plants in the main periods of time.

Such a diagram can only represent the present state of our imperfect knowledge; any day discoveries may extend the line of any group up or down in the series, or may connect the groups together.

It becomes evident that so early as the Palæozoic there are nearly as many types represented as in the present day, and that in fact everything, up to the higher Gymnosperms, was well developed (for it is hard indeed to prove that Cordaites is less highly organized than some of the present Gymnosperm types), but flowering plants and also the true cycads are wanting, as well as the intermediate Mesozoic Bennettitales. The peculiar groups of the period were the Pteridosperm series, connecting links between fern and cycad, and the Sphenophyllums, connecting in some measure the Lycopods and Calamites. With them some of the still living groups of ferns, Lycopods, and Equisetaceæ were flourishing, though all the species differed from those now extant. This shows us how very far from the beginning our earliest information is, for already in the Palæozoic we have a flora as diversified as that now living, though with more primitive characters.

fig122

Fig. 122.:Diagram showing the relative distribution of the main groups of plants through the geological eras. The dotted lines connecting the groups and those in the pre-Carboniferous are entirely theoretical, and merely indicate the conclusions reached at present. The size of the surface of each group roughly indicates the part it played in the flora of each period. Those with dotted surface bore seeds, the others spores.

In Mesozoic times the most striking group is that of the Cycads and Bennettitales, the latter branch suggesting a direct connection between the fern-cycad series and the flowering plants. This view, so recently published and upheld by various eminent botanists, is fast gaining ground. Indeed, so popular has it become among the specialists that there is a danger of overlooking the real difficulties of the case. The morphological leap from the leaves and stems of cycads to those of the flowering plants seems a much more serious matter to presuppose than is at present recognized.

As is indicated in the diagram, the groups do not appear isolated by great unbridged gaps, as they did even twenty years ago. By means of the fossils either direct connections or probable lines of connection are discovered which link up the series of families. At present the greatest gap now lies hedging in the Moss family, and, as was mentioned, fossil botany cannot as yet throw much light on that problem owing to the lack of fossil mosses.

This glimpse into the past suggests a prophecy for the future. Evolution having proceeded steadily for such vast periods is not likely to stop at the stage reached by the plants of to-day. What will be the main line of advance of the plants of the future, and how will they differ from those of the present?

We have seen in the past how the differentiation of size in the spores resulted in sex, and in the higher plants in the modifications along widely different lines of the male and female; how the large spore (female) became enclosed in protecting tissues, which finally led up to true seeds, while the male being so temporary had no such elaboration. As the seed advances it becomes more and more complex, and when we reach still higher plants further surrounding tissues are pressed into its service and it becomes enclosed in the carpel of the highest flowering plants. After that the seed itself has fewer general duties, and instead of those of the Gymnosperms with large endosperms collecting food before the embryo appears, small ovules suffice, which only develop after fertilization is assured. The various families of flowering plants have gone further, and the whole complex series of bracts and fertile parts which make up a flower is adapted to ensure the crossing of male and female of different individuals. The complex mechanisms which seem adapted for “cross fertilization” are innumerable, and are found in the highest groups of the flowering plants. But some have gone beyond the stage when the individual flowers had each its device, and accomplished its seed-bearing independently of the other flowers on the same branch. These have a combination of many flowers crowded together into one community, in which there is specialization of different flowers for different duties. In such a composite flower, the Daisy for example, some are large petalled and brightly coloured to attract the pollen-carrying insects, some bear the male organs only, and others the female or seed-producing. Here, then, in the most advanced type of flowering plant we get back again to the separation of the sexes in separate flowers; but these flowers are combined in an organized community much more complex than the cones of the Gymnosperms, for example, where the sexes are separate on a lower plane of development.

It seems possible that an important group, if not the dominant group, of flowering plants in the future will be so organized that the individual flowers are very simple, with fewer parts than those of to-day, but that they will be combined in communities of highly specialized individuals in each flower head or cluster.

As well as this, in other species the minute structure of the vital organs may show a development in a direction contrary to what has hitherto seemed advance. Until recently flowers and their organs have appeared to us to be specialized in the more advanced groups on such lines as encourage “cross fertilization”. In “cross fertilization”, in fact, has appeared to lie the secret of the strength and advance of the races of plants. But modern cytologists have found that many of the plants long believed to depend on cross fertilization are either self-fertilized or not fertilized at all! They have passed through the period when their complex structures for ensuring cross fertilization were used, and though they retain these external structures they have taken to a simpler method of seed production, and in some cases have even dispensed with fertilization of the egg cell altogether. The female vitality increased, the male becomes superfluous. It is simpler and more direct to breed with only one sex, or to use the pollen of the same individual. Many flowers are doing this which until recently had not been suspected of it. We cannot yet tell whether it will work successfully for centuries to come or is an indication of “race senility”.

Whether in the epochs to come flowering plants will continue to hold the dominant position which they now do is an interesting theoretical problem. Flowers were evolved in correlation with insect pollination. One can conceive of a future, when all the earth is under dominion of man, in which fruits will be sterilized for man’s use, as the banana is now, and seed formation largely replaced by gardeners’ “cuttings”.

In those plants which are now living where the complex mechanisms for cross-fertilization have been superseded by simple self-fertilization, the external parts of the more elaborate method are still produced, though they are apparently futile. In the future these vestigial organs will be discarded, or developed in a more rudimentary form (for it is remarkable how organs that were once used by the race reappear in members of it that have long outgrown their use), and the morphology of the flower will be greatly simplified.

Thus we can foresee on both sides much simplified individual flowers:in the one group the reduced individuals associating together in communities the members of which are highly specialized, and in the other the solitary flowers becoming less elaborate and conspicuous, as they no longer need the assistance of insects (the cleistogamic flowers of the Violet, for example, even in the present day bend toward the earth, and lack all the bright attractiveness of ordinary flowers), and perhaps finally developing underground, where the seeds could directly germinate.

In the vegetative organs less change is to be expected, the examples from the past lead us to foresee no great difference in size or general organization of the essential parts, though the internal anatomy has varied, and probably will vary, greatly with the whole evolution of the plant.

But one more point and we must have done. Why do plants evolve at all? Why did they do so through the geological ages of the past, and why should we expect them to do so in the future? The answer to this question must be less assured than it might have been even twenty years ago, when the magnetism of Darwin’s discoveries and elucidations seemed to obsess his disciples. “Response to environment” is undoubtedly a potent factor in the course of evolution, but it is not the cause of it. There seems to be something inherent in life, something apparently (though that may be due to our incomplete powers of observation) apart from observable factors of environment which causes slight spontaneous changes, mutations, and some individuals of a species will suddenly develop in a new direction in one or other of their parts. If, then, this places them in a superior position as regards their environment or neighbours, it persists, but if not, those individuals die out. The work of a special branch of modern botany seems clearly to indicate the great importance of this seemingly inexplicable spontaneity of life. In environment alone the thoughtful student of the present cannot find incentive enough for the great changes and advances made by organisms in the course of the world’s history. The climate and purely physical conditions of the Coal Measure period were probably but little different from those in some parts of the world to-day, but the plants themselves have fundamentally changed. True, their effect upon each other must be taken into account, but this is a less active factor with plants than with men, for we can imagine nothing equivalent to citizenship, society, and education in the plant communities, which are so vital in human development.

It seems to have been proved that plants and animals may, at certain unknown intervals, “mutate”; and mutation is a fine word to express our recent view of one of the essential factors in evolution. But it is a cloak for an ignorance avowedly less mitigated than when we thought to have found a complete explanation of the causes of evolution in “environment”.

In a sketch such as the present, outlines alone are possible, detail cannot be elaborated. If it has suggested enough of atmosphere to show the vastness of the landscape spreading out before our eyes back into the past and on into the future, the task has been accomplished. There are many detailed volumes which follow out one or other special line of enquiry along the highroads and by-ways of this long traverse in creation. If the bird’s-eye view of the country given in this book entices some to foot it yard by yard under the guidance of specialists for each district, it will have done its part. While to those who will make no intimate acquaintance with so far off a land it presents a short account by a traveller, so that they may know something of the main features and a little of the romance of the fossil world.

Ancient Plants – Stages in Plant Evolution

CHAPTER V
STAGES IN PLANT EVOLUTION

To attempt any discussion of the causes of evolution is far beyond the scope of the present work. At present we must accept life as we find it, endowed with an endless capacity for change and a continuous impulse to advance. We can but study in some degree the course taken by its changes.

From the most primitive beginnings of the earliest periods, enormous advance had been made before we have any detailed records of the forms. Yet there remain in the world of to-day numerous places where the types with the simplest structure can still flourish, and successfully compete with higher forms. Many places which, from the point of view of the higher plants, are undesirable, are well suited to the lower. Such places, for example, as the sea, and on land the small nooks and crannies where water drops collect, which are useless for the higher plants, suffice for the minute forms. In some cases the lower plants may grow in such masses together as to capture a district and keep the higher plants from it. Equisetum (the horsetail) does this by means of an extensive system of underground rhizomes which give the plant a very strong hold on a piece of land which favors it, so that the flowering plants may be quite kept from growing there.

In such places, by a variety of means, plants are now flourishing on the earth which represent practically all the main stages of development of plant life as a whole. It is to the study of the simpler of the living forms that we owe most of our conceptions of the course taken by evolution. Had we to depend on fossil evidence alone, we should be in almost complete ignorance of the earliest types of vegetation and all the simpler cohorts of plants, because their minute size and very delicate structure have always rendered them unsuitable for preservation in stone. At the same time, had we none of the knowledge of the numerous fossil forms which we now possess, there would be great gaps in the series which no study of living forms could supply. It is only by a study and comparison of both living and fossil plants of all kinds and from beds of all ages that we can get any true conception of the whole scheme of plant life.

Grouping together all the main families of plants at present known to us to exist or to have existed, we get the following series::

Group. Common examples of typical families in the group.

Thallophyta

Algæ Seaweeds.

Fungi Moulds and toadstools.

Bryophyta

Hepaticæ Liverworts.

Musci Mosses.

Pteridophyta

Equisetales Horsetails.

Sphenophyllales* fossil only, Sphenophyllum.

Lycopodales Club-moss.

Filicales Bracken fern.

Pteridospermæ

Lyginodendræ* fossil only, Sphenopteris.

Gymnosperms

Cycadales Cycads.

Bennettitales* fossil only, Bennettites.

Ginkgoales Maidenhair Tree.

Cordaitales* fossil only, Cordaites.

Coniferales Pine, Yew.

Gnetales Welwitschia.

Angiosperms

Monocotyledons Lily, Palm, Grass.

Dicotyledons Rose, Oak, Daisy.

In this table the different groups have not a strictly equivalent scientific value, but each of those in the second column represents a large and well-defined series of primary importance, whose members could not possibly be included along with any of the other groups.

Those marked with an asterisk are known only as fossils, and it will be seen that of the seventeen groups, so many as four are known only in the fossil state. This indicates, however, but a part of their importance, for in nearly every other group are many families or genera which are only known as fossils, though there are living representatives of the group as a whole.

In this table the individual families are not mentioned, because for the present we need only the main outline of classification to illustrate the principal facts about the course of evolution. As the table is given, the simplest families come first, the succeeding ones gradually increasing in complexity till the last group represents the most advanced type with which we are acquainted, and the one which is the dominant group of the present day.

This must not be taken as a suggestion that the members of this series have evolved directly one from the other in the order in which they stand in the table. That is indeed far from the case, and the relations between the groups are highly complex.

It must be remarked here that it is often difficult, even impossible, to decide which are the most highly evolved members of any group of plants. Each individual of the higher families is a very complicated organism consisting of many parts, each of which has evolved more or less independently of the others in response to some special quality of the surroundings. For instance, one plant may require, and therefore evolve, a very complex and well-developed water-carriage system while retaining a simple type of flower; another may grow where the water problem does not trouble it, but where it needs to develop special methods for getting its ovules pollinated; and so on, in infinite variety. As a result of this, in almost all plants we have some organs highly evolved and specialized, and others still in a primitive or relatively primitive condition. It is only possible to determine the relative positions of plants on the scale of development by making an average conclusion from the study of the details of all their parts. This, however, is beset with difficulties, and in most cases the scientist, weighed by personal inclinations, arbitrarily decides on one or other character to which he pays much attention as a criterion, while another scientist tends to lay stress on different characters which may point in another direction.

In no group is this better illustrated than among the Coniferæ, where the relative arrangement of the different families included in it is still very uncertain, and where the observations of different workers, each dealing mainly with different characters in the plants, tend to contradict each other.

This, however, as a byword. Notwithstanding these difficulties, which it would be unfair to ignore, the main scheme of evolution stands out clearly before the scientist of to-day, and his views are largely supported by many important facts from both fossil and living plants.

Very strong evidence points to the conclusion that the most primitive plants of early time were, like the simplest plants of to-day, water dwellers. Whether in fresh water or the sea is an undecided point, though opinion seems to incline in general to the view that the sea was the first home of plant life. It can, however, be equally well, and perhaps even more successfully argued, that the freshwater lakes and streams were the homes of the first families from which the higher plants have gradually been evolved.

For this there is no direct evidence in the rocks, for the minute forms of the single soft cells assumed by the most primitive types were just such as one could not expect to be successfully fossilized. Hence the earliest stages must be deduced from a comparative study of the simplest plants now living. Fortunately there is much material for this in the numerous waters of the earth, where swarms of minute types in many stages of complexity are to be found.

The simplest type of plants now living, which appears to be capable of evolution on lines which might have led to the higher plants, is that found in various members of the group of the Protococcoideæ among the Algæ. The claim of bacteria and other primitive organisms of various kinds to the absolute priority of existence is one which is entirely beyond the scope of a book dealing with fossil plants. The early evolution of the simple types of the Protococcoideæ is also somewhat beyond its scope, but as they appear to lie on the most direct “line of descent” of the majority of the higher plants it cannot be entirely ignored. From the simpler groups of the green Algæ other types have specialized and advanced along various directions, but among them there seems an inherent limitation, and none but the protococcoid forms seem to indicate the possibility of really high development.

fig17

Fig. 17.:A Protococcoid Plant consisting of one cell

p, Protoplasm; n, nucleus; g, coloring body or chloroplast; w, cell wall.

In a few words, a typical example of one of the simple Protococcoideæ may be described as consisting of a mass of protoplasm in which lie a recognizable nucleus and a green coloring body or chloroplast, with a cell wall or skin surrounding these vital structures, a cell wall that may at times be dispensed with or unusually thickened according as the need arises. This plant is represented in fig. 17 in a somewhat diagrammatic form.

In such a case the whole plant consists of one single cell, living surrounded by the water, which supplies it with the necessary food materials, and also protects it from drying up and from immediate contact with any hard or injurious object. When these plants propagate they divide into four parts, each one similar to the original cell, which all remain together within the main cell wall for a short time before they separate.

If now we imagine that the four cells do not separate, but remain together permanently, we can see the possibility of a beginning of specialization in the different parts of the cell. The single living cell is equally acted on from all sides, and in itself it must perform all the life functions; but where four lie together, each of the four cells is no longer equally acted on from all sides. This shows clearly in the diagram of a divided cell given in fig. 18. Here it is obvious that one side of each of the four cells, viz. that named a in the diagram, is on the outside and in direct contact with the water and external things; but walls b and c touch only the corresponding walls of the neighboring cells. Through walls b and c no food and water can enter directly, but at the same time they are protected from injury and external stimulus. Hence, in this group of four cells there is a slight differentiation of the sides of the cells. If now we imagine that each of the four cells, still remaining in contact, divides once more into four members, each of which reaches mature size while all remain together, then we have a group of sixteen cells, some of which will be entirely inside, and some of which will have walls exposed to the environment.

fig18

Fig. 18.:Diagram of Protococcoid Cell divided into four daughter cells. Walls a are external, and walls b and c in contact with each other.

If the cells of the group all divide again, in the manner shown the mass will become more than one cell thick, and the inner cells will be more completely differentiated, for they will be entirely cut off from the outside and all direct contact with water and food materials, and will depend on what the outer cells transmit to them. The outer cells will become specialized for protection and also for the absorption of the water and salts and air for the whole mass. From such a plastic group of green cells it is probable that the higher and increasingly complex forms of plants have evolved. There are still living plants which correspond with the groups of four, sixteen, &c., cells just now theoretically stipulated.

fig19

Fig. 19.:A, Details of Part of the Tissues in a Stem of a Flowering Plant. B, Diagram of the Whole Arrangement of Cross Section of a Stem: e, Outer protecting skin; g, green cells; s, thick-walled strengthening cells; p, general ground tissue cells. V, Groups of special conducting tissues: x, vessels for water carriage; px, first formed of the water vessels; c, growing cells to add to the tissues; b, food-conducting cells; ss, strengthening cells.

The higher plants of to-day all consist of very large numbers of cells forming tissues of different kinds, each of which is specialized more or less, some very elaborately, for the performance of certain functions of importance for the plant body as a whole. With the increase in the number of cells forming the solid plant body, the number of those living wholly cut off from the outside becomes increasingly great in comparison with those forming the external layer. Some idea of the complexity and differentiation of this cell mass is given in fig. 19, A, which shows the relative sizes and shapes of the cells composing a small part of the stem of a common flowering plant. The complete section would be circular and the groups V would be repeated round it symmetrically, and the whole would be enclosed by an unbroken layer of the cells marked e, as in the diagram B.

fig20

Fig. 20.:Conducting Cells and Surrounding Tissue seen in fig. 19, A, cut lengthways. px, First formed vessels for water conduction; x, larger vessel; b, food-conducting cells; ss, strengthening cells; p, general ground tissue.

In the tissues of the higher plants the most important feature is the complex system of conducting tissues, shown in the young condition in V in fig. 19, A. In them the food and water conducting elements are very much elongated and highly specialized cells, which run between the others much like a system of pipes in the brickwork of a house. These cells are shown cut longitudinally in fig. 20, where they are lettered to correspond with the cells in fig. 19, A, with which they should be compared. In such a view the great difference between the highly specialized cells x, px, b, &c., and those of the main mass of ground tissue p becomes apparent.

Even in the comparatively simply organized groups of the Equisetales and Lycopodiales the differentiation of tissues is complete. In the mosses, and still more in the liverworts, it is rudimentary; but they grow in very damp situations, where the conduction of water and the protection from too much drying is not a difficult problem for them. As plants grow higher into the air, or inhabit drier situations, the need of specialization of tissues becomes increasingly great, for they are increasingly liable to be dried, and therefore need a better flow of water and a more perfect protective coat.

It is needless to point out how the individual cells of a plant, such as that figured in figs. 19 and 20, have specialized away from the simple type of the protococcoid cell in their mature form. In the young growing parts of a plant, however, they are essentially like protococcoid cells of squarish outline, fitting closely to each other to make a solid mass, from which the individual types will differentiate later and take on the form suitable for the special part they have to play in the economy of the whole plant.

To trace the specialization not only of the tissues but of the various parts of the whole plant which have become elaborate organs, such as leaves, stems, and flowers, is a task quite beyond the present work to attempt. From the illustrations given of tissue structure from plants at the two ends of the series much can be imagined of the inevitable intermediate stages in tissue evolution.

As regards the elaboration of organs, and particularly of the reproductive organs, details will be found throughout the book. In judging of the place of any plant in the scale of evolution it is to the reproductive organs that we look for the principal criteria, for the reproductive organs tend to be influenced less by their physical surroundings than the vegetative organs, and are therefore truer guides to natural relationships.

In the essential cells of the reproductive organs, viz. the egg cell and the male cell, we get the most primitively organized cells in the plant body. In the simpler families both male and female cells return to the condition of a free-swimming protococcoid cell, and in all but the highest families the male cell requires a liquid environment, in which it swims to the egg cell. In the higher families the necessary water is provided within the structure of the seed, and the male cell does not swim, a naked, solitary cell, out into the wide world, as it does in all the families up to and including the Filicales. In the Coniferæ and Angiosperms the male cell does not swim, but is passive (or largely so), and is brought to the egg cell. One might almost say that the whole evolution of the complex structures found in fruiting cones and flowers is a result of the need of protection of the delicate, simple reproductive cells and the embryonic tissues resulting from their fusion. The lower plants scatter these delicate cells broadcast in enormous numbers, the higher plants protect each single egg cell by an elaborate series of tissues, and actually bring the male cell to it without ever allowing either of them to be exposed.

It must be assumed that the reader possesses a general acquaintance with the living families tabulated on p. 44; those of the fossil groups will be given in some detail in succeeding chapters which deal with the histories of the various families. It is premature to attempt any general discussion of the evolution of the various groups till all have been studied, so that this will be reserved for the concluding chapters.