Fig. 17.—A Protococcoid Plant consisting of one cell
p, Protoplasm;n, nucleus;g, colouring 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 colouring 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 infig. 17in 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 possibilityof 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 infig. 18. Here it is obvious that one side of each of the four cells, viz. that namedain the diagram, is on the outside and in direct contact with the water and external things; but wallsbandctouch only the corresponding walls of the neighbouring cells. Through wallsbandcno 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.
Fig. 18.—Diagram of Protococcoid Cell divided into four daughter cells. Wallsaare external, and wallsbandcin 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.
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 theoutside becomes increasingly great in comparison with those forming the external layer. Some idea of the complexity and differentiation of this cell mass is given infig. 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 groupsVwould be repeated round it symmetrically, and the whole would be enclosed by an unbroken layer of the cells markede, as in the diagramB.
Fig. 20.—Conducting Cells and Surrounding Tissue seen infig. 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 inVinfig. 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 infig. 20, where they are lettered to correspond with the cells infig. 19,A, with which they should becompared. In such a view the great difference between the highly specialized cellsx,px,b, &c., and those of the main mass of ground tissuepbecomes 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.19and20, 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 physicalsurroundings 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 itswimsto 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 onp. 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.
The individual plants of the Coal Measure period differed entirely from those now living; they were more than merely distinct species, for in the main even the families were largely different from the present ones. Nevertheless, when we come to examine the minute anatomy of the fossils, and the cells of which they are composed, we find that between the living and the fossil cell types the closest similarity exists.
From the earliest times of which we have any knowledge the elements of the plant body have been the same, though the types of structures which they built have varied in plan. Individualcellsof nearly every type from the Coal Measure period can be identically matched with those of to-day. In the way the walls thickened, in the shapes of the wood, strengthening or epidermal cells, in the form of the various tissues adapted to specific purposes, there is a unity of organization which it is reasonable to suppose depends on the fundamental qualities inherent in plant life.
This will be illustrated best, perhaps, by tabulating the chief modifications of cells which are found in plant tissues. The illustrations of these types in the following table are taken from living plants, because from them figures of more diagrammatic clearness can be made, and the salient characters of the cells more easily recognized. Comparison of these typical cells with those illustrated from the fossil plants reveals their identity in essential structure, and most of them will be found in the photos of fossils in these pages, though they are better recognized in the actual fossils themselves.
Epidermal.
Fig. 21
Epidermis.—Protecting layer or skin. Cells with outer wall thickened in many cases (fig. 21,aandb). Compare fossil epidermis infig. 34,e.
Fig. 22
Hairs.—Extensions of epidermis cells. Single cells, or complex, asfig. 22,h, whereeis epidermis andpparenchyma. Compare fossil hairs in figs.79and120.
Fig. 23
Stomates.—Breathing pores in the epidermis. Seen in surface view as two-lipped structures (fig. 23).s, Stomates;e, epidermis cells. Compare fossil stomates infig. 8.
Ground Tissue
Fig. 24
Parenchyma.—Simple soft cells, either closely packed, as infig. 24, or with air spaces between them. Compare78,B, for fossil.
Fig. 25
Palisade.—Elongated, closely packed cells,p, chiefly in leaves, lying below the epidermis,e,fig. 25. Comparefig. 34,p, for fossil palisade.
Fig. 26
Endodermis.—Cells with specially thickened walls,en, lying as sheath between the parenchyma,c, of ground tissue, and the vascular tissue,s,fig. 26. Comparefig. 108for fossil endodermis.
Fig. 27
Latex cells.—Large, often much elongated cells,m, lying in the parenchyma,p,fig. 27, which are packed with contents. Comparefig. 107,s.
Fig. 28
Sclerenchyma.—Thick-walled cells among parenchyma for strengthening,fig. 28. Comparefig. 34,s.
Fig. 29
Cork.—Layers of cells replacing the epidermis in old stems. Outer cells,o, crushed;k, closely packed cork cells; stone cells,s,fig. 29. Comparefig. 95,k.
Cork cambium.—Narrow, actively dividing cells,cinfig. 29, giving rise to new cork cells in consecutive rows.
Fig. 30
Tracheides.—Specially thickened cells in the parenchyma, usually for water storage,t,fig. 30. Comparefig. 95,t.
Vascular Tissue
Fig. 31
Wood.—Protoxylem, tracheids and vessels, long, narrow elements, with spiral or ring-like thickenings,s1ands2,fig. 31. Comparefig. 81,A,px, for fossil.
Metaxylem, long elements, tracheids and vessels. Some with narrow pits, astinfig. 31; others with various kinds of pits. In transverse section seen infig. 33, w, fossil infig. 114,w.
Wood parenchyma.—Soft cells associated with the wood,pinfig. 31. Fossil infig. 81,B,p.
Wood sclerenchyma.—Hard thickened cells in the wood.
Fig. 32
Bast.—Sieve tubes, long cells which carry foodstuffs, cross walls pitted like sieves,s,fig. 32. In transverse section infig. 33.
Companion cells, narrow cells with rich proteid contents,c,fig. 32. In transverse section atc,fig. 33.
Bast parenchyma.—Soft unspecialized cells mixed with the sieve tubes,p,fig. 32.
Bast fibres.—Thick-walled sclerenchymatous cells mixed with, or outside, the soft bast.
Fig. 33
Cambium.—Narrow cells, like those of the cork cambium, which lie between the wood and bast, and give rise to new tissues of each kind,cb,fig. 33. Comparefig. 114, fossil.
There are, of course, many minor varieties of cells, but these illustrate all the main types.
Among the early fossils, however, one type of wood cell and one type of bast cell, so far as we know, are not present. These cells are the truevesselsof the wood of flowering plants, and the long bast cells with their companion proteid cells. The figure of a metaxylem wood cell, shown infig. 31,t, shows the more primitive type of wood cell, which has an oblique cross wall. This type of wood cell is found in all the fossil trees, and all the living plants except the flowering plants. The vessel type, which is that in the big wood vessels of the flowering plants, and has no cross wall, is seen infig. 20,x.
The similarity between the living cells and those of the Coal Measure fossils is sufficiently illustrated to need no further comment. This similarity is an extremely helpful point when we come to an interpretation of the fossils. In living plants we can study the physiology of the various kinds of cells, and can deduce from experiment exactly the part they play in the economy of the whole plant. From a study of the tissues in any plant structure we know what function it performed, and can very often estimate the nature of the surrounding conditions under which the plant was growing. To take a single example, the palisade tissue, illustrated infig. 25,p, in living plants always contains green colouring matter, and lies just below the epidermis, usually of leaves, but sometimes also of green stems. These cells do most of the starch manufacture for the plant, and are found best developed when exposed to a good light. In very shady places the leaves seldom have this type of cell. Now, when cells just like these are found in fossils (as is illustrated infig. 34), we can assume all the physiological facts mentioned above, and rest assured that that leaf was growing under normal conditions of light and was actively engaged in starch-building when it was alive. From the physiological standpoint the fossil leaf is entirely the same as a normal living one.
Fig. 34.—From a Photo of a Fossil Lea
e, Epidermis;p, palisade cells;pr, soft parenchyma cells (poorly preserved);s, sclerenchyma above the vascular bundle.
From the morphological standpoint, also, the features of the plant body from the Coal Measure period fall into the same divisions as those of the present. Roots, stems, leaves, and reproductive organs, the essentially distinct parts of a plant, are to be found in a form entirely recognizable, or sufficiently like that now in vogue to be interpreted without great difficulty. In the detailed structure of the reproductive organs more changes have taken place than in any others, both in internal organization and external appearance.
Already, in the Early Palæozoic period, the distinction between leaves, stems, roots, and reproductive organs was as clearly marked as it is to-day, and, judging by their structure, they must each have performed the physiological functions they now do. Roots have changed least in the course of time, probably because, in the earth, they live under comparatively uniform conditions in whatever period of the world’s history they are growing. Naturally, between the roots of different species there are slight differences; but the likeness between fern roots from the Palæozoic and from a living fern is absolutely complete. This is illustrated infig. 35, which shows themicroscopic structure of the two roots when cut in transverse direction. The various tissues will be recognized as coming into the table onp. 54, so that both in the details of individual cells and in the general arrangement of the cell groups or tissues the roots of these fossil and living ferns agree.
Fig. 35.—A, Root of Living Fern.B, Root of Palæozoic Fossil Fern.c, Cortex;px, protoxylem in two groups;m, metaxylem;s, space in fossil due to decay of soft cells.
Among stems there has been at all periods more variety than among the roots of the corresponding plants, and in the following chapter, when the differences between living and fossil plants will be considered, there will be several important structures to notice. Nevertheless, there are very many characters in which the stems from such widely different epochs agree. The plants in the palæozoic forests were of many kinds, and among them were those with weak trailing stems which climbed over and supported themselves on other plants, and also tall, sturdy shafts of woody trees, many of which were covered with a corky bark. Leaves were attached to the stems, either directly, as in the case of some living plants, or by leaf stalks. In external appearance and in general function the stems then were as stems are now. In the details of the individual cells also the likeness is complete; it is in the grouping of the cells, the anatomy of the tissues, that the important differences lie. It hasbeen remarked already that increase in complexity of the plant form usually goes with an increase in complexity of the cells and variety of the tissues. The general ground tissue in nearly all plants is very similar; it is principally in the vascular system that the advance and variety lie.
Plant anatomists lay particular stress on the vascular system, which, in comparison with animal anatomy, holds an even more important position than does the skeleton. To understand the essential characters of stems, both living and fossil, and to appreciate their points of likeness or difference, it is necessary to have some knowledge of the general facts of anatomy; hence the main points on which stress is laid will be given now in brief outline.
Leaving aside consideration of the more rudimentary and less defined structure of the algæ and mosses, all plants may be said to possess a “vascular system”. This is typically composed of elongated wood (or xylem) with accessory cells (seep. 57, table), and bast (phloem), also with accessory cells. These specialized conducting elements lie in the ground tissue, and in nearly all cases are cut off from direct contact with it by a definite sheath, called the endodermis (seep. 55,fig. 26). Very often there are also groups or rings of hard thick-walled cells associated with the vascular tissues, which protect them and play an important part in the consolidation of the whole stem.
Fig. 36.—Diagram of Simplest Arrangement of Complete Stele in a Stem
W, Central solid wood;P, ring of bast;E, enclosing sheath of endodermis;C, ground tissue or cortex.
The simplest, and probably evolutionally the mostprimitive form which is taken by the vascular tissues, is that of a single central strand, with the wood in the middle, the bast round it, and a circular endodermis enclosing all, as infig. 36, which shows a diagram of this arrangement. Such a mass of wood and bast surrounded by an endodermis, is technically known as astele, a very convenient term which is much used by anatomists. In its simplest form (as infig. 36) it is called aprotostele, and is to be found in both living and fossil plants. A number of plants which get more complex steles later on, have protosteles in the early stages of their development, as inPteris auritafor example, a species allied to the bracken fern, which has a hollow ring stele when mature.
Fig. 37.—Diagram of a Stele with a few Cells of Pithpin the Middle of the Wood. Lettering as infig. 36
Fig. 38.—Diagram showing Extensive Pithpin the Wood. Lettering as infig. 36
The next type of stele is quite similar to the protostele, but with the addition of a few large unspecialized cells in the middle of the wood (p,fig. 37); these are the commencement of the hollowing process which goes on in the wood, resulting later in the formation of a considerable pith, as is seen infig. 38, where the wood is now a hollow cylinder, as the phloem has been from the first. When this is the case, a secondsheath or endodermis generally develops on the inner side of the wood, outside the pith, and cuts the vascular tissues off from the inner parenchyma. A further step is the development of an inner cylinder of bast so that the vascular ring is completely double, with endodermis on both sides of the cylinder, as is seen infig. 39.
Fig. 39.—A Cylindrical Stele, withe, inner endodermis, andph, inner phloem;W, wood;P, outer phloem;E, outer endodermis.L, part of the stele going out to supply a large leaf, thus breaking what would otherwise appear as a closed ring stele
In all these cases there is but one strand or cylinder, of vascular tissue in the stem, but one stele, and this type of anatomy is known as themonostelicor single-steled type.
Fig. 40.—A Ring Stele apparently broken up into a Number of Protosteles by many Leaf Gaps
When from the double cylinder just described a strand of tissue goes off to supply a large leaf, a considerable part of the stele goes out and breaks the ring. This is shown infig. 39, whereLis the part of the stele going to a leaf, and the rest the broken central cylinder. When the stem is short, and leaves grow thickly so that bundles are constantly going out from the main cylinder, this gets permanently broken, and its appearance when cut across at any given point is that of a group of several steles arranged in a ring, each separate stele being like the simple protostele in its structure. Seefig. 40. This type of stem has long been known aspolystelic(i.e.many-steled), andit is still a convenient term to describe it by. There has been much theoretical discussion about the true meaning of such a “polystelic” stem, which cannot be entered into here; it may be noted, however, that the various strands of the broken ring join up and form a meshwork when we consider the stem as a whole, it is only in a single section that they appear as quite independent protosteles. Nevertheless, as we generally consider the anatomy of stems in terms of single sections, and as the descriptive word “polystelic” is a very convenient and widely understood term, it will be used throughout the book when speaking of this type of stem anatomy.
Such a type as this, shown infig. 40, is already complex, but it often happens that the steles branch and divide still further, until there is a highly complicated and sometimes bewildering system of vascular strands running through the ground tissue in many directions, but cut off from it by their protective endodermal sheaths. Such complex systems are to be found both in living and fossil plants, more especially in many of the larger ferns (seefig. 88).
Higher plants in general, however, and in particular flowering plants, do not have a polystelic vascular arrangement, but a specialized type of monostele.
Fig. 41.—Monostele in which the Central Pith is Star-shaped, and the Wood breaking up into Separate Groups
p, Pith;W, wood;P, phloem;E, endodermis;C, cortex.
Referring again tofig. 37as a starting-point, imagine the pith in the centre to spread in a star-shaped form till the points of the star touched the edges of the ring, and thus to break the wood ring into groups. A stage in this process (which is not yet completed) is shown infig. 41, while infig. 42the wood and bast groupsare entirely distinct. In the flowering plants the cells of the endodermis are frequently poorly characterized, and the pith cells resemble those of the cortical ground tissue, so that the separate groups of wood and bast (usually known as “vascular bundles”, in distinction from the “steles” offig. 40) appear to lie independently in the ground tissue. These strands, however, must not be confused with steles, they are only fragments of the single apparently broken up stele which runs in the stem.
Fig. 42.—Monostele in which the Pith has invaded all the Tissues as far as the Endodermis, and broken the Wood and Phloem up into Separate Bundles. These are usually called “vascular bundles” in the flowering plants
Fig. 43.—Showing actively growing Zonec(Cambium) in the Vascular Bundles, and joining across the ground tissue between them
The vascular bundle, of all except the Monocotyledons, has a potentiality for continued growth and expansion which places it far above the stele in value for a plant of long life and considerable growth. The cells lying between the wood and the bast, the soft parenchyma cells always accompanying such tissues, retain their vitality and continue to divide with great regularity, and to give rise to a continuous succession of new cells of wood on the one side and bast on the other; seefig. 33,c,b. In this way the primary, distinct vascular bundles are joined by a ring of wood, seefig. 43, to which are added further rings every season, till the mass of wood becomes a strong solid shaft. This ever-recurring activity of the cambium gives rise to what are known as “annual rings” in stems, seefig. 44, in which the woodshows both primary distinct groups in the centre, and the rings of growth of later years.
Cambium with this power of long-continued activity is found in nearly all the higher plants of to-day (except the Monocotyledons), but in the fern and lycopod groups it is in abeyance. Certain cases from nearly every family of the Pteridophytes are known, where some slight development of cambium with its secondary thickening takes place, but in the groups below the Gymnosperms cambium has almost no part to play. On the other hand, so far back as the Carboniferous period, the masses of wood in the Pteridophyte trees were formed by cambium in just the same way as they are now in the higher forms. Its presence was almost universal at that time in the lower groups where to-day there are hardly any traces of it to be found.
Fig. 44.—Stem with Solid Cylinder of Wood developed from the Cambium, showing three “annual rings”. In the centre may still be seen the separate groups of the wood of the primary “vascular bundles”
It will be seen from this short outline of the vascular system of plants, that there is much variety possible from modifications of the fundamental protostele. It is also to be noted that the plants of the Coal Measures had already evolved all the main varieties of steles which are known to us even now,[6]and that the development of secondary thickening was very widespread. In several cases the complexity of type exceeds that ofmodern plants (seeChap. VII), and there are to be found vascular arrangements no longer extant.
When we turn to theReproductive Organs, we find that the points of likeness between the living and the fossil forms are not so numerous or so direct as they are in the case of the vegetative system.
Fig. 45.—Fern Sporangia
A, fossil;B, living.
As has been indicated, the families of plants typical of the Coal Measures were not those which are the most prominent to-day, but belonged to the lower series of Pteridophytes. In their simpler forms the fructifications then and now resemble each other very closely, but in the more elaborate developments the points of variety are more striking, so that they will be dealt with in the following chapter. Cases of likeness are seen in the sporangia of ferns, some of which appear to have been practically identical with those now living. This is illustrated infig. 45, which shows the outline of the cells of the sporangia of living and fossil side by side.
Fig. 46.—A, Living Lycopod cone;B,Lepidodendron(fossil) cone.a, Axis;s, scale;S, sporangium with spores. One side of a longitudinal section
In the general structure also of the cones of the simpler types ofLepidodendron(fossil, seefrontispiece) there is a close agreement with the living Lycopods,though as regards size and output of spores there was a considerable difference in favour of the fossils. The plan of each is that round the axis of the cone simple scales are arranged, on each of which, on its upper side, is seated a large sporangium bearing numerous spores all of one kind (seefig. 46).
Equally similar are the cones of the living Equisetum and some of the simple members of the fossil family Calamiteæ, but the more interesting cases are those where differences of an important morphological nature are to be seen.
As regards the second[7]generation there is some very important evidence, from extremely young stages, which has recently been given to the world. In a fern sporangiumgerminating sporeswere fossilized so as to show the first divisions of the spore cell. These seem to be identical with the first divisions of some recent ferns (seefig. 47). This is not only of interest as showing the close similarity in detail between plants of such widely different ages, but is a remarkable case of delicate preservation of soft and most perishable structures in the “coal balls”.