CHAPTER XXXVIII.

THE ORGANIZATION OF THE PLANT.

I. Organization of Plant Members.[37]

=689.= It is now generally conceded that the earliest plants to appear in the world were very simple in form and structure. Perhaps the earliest were mere bits of naked protoplasm, not essentially different from early animal life. The simplest ones which are clearly recognized as plants are found among the lower algæ and fungi. These are single cells of very minute size, roundish, oval, or oblong, existing during their growing period in water or in a very moist substratum or atmosphere. Examples are found in the red snow plant (_Sphærella nivalis_), the Pleurococcus, the bacteria; and among small colonies of these simple organisms (Pandorina) or the thread-like forms (Spirogyra, Œdogonium, etc.). It is evident that some of the life relations of such very simple organisms are very easily obtained—that is, the adjustment to environment is not difficult. All of the living substance is very closely surrounded by food material in solution. These food solutions are easily absorbed. Because of the minute size of the protoplasts and of the plant body, they do not have to solve problems of transport of food to distant parts of the body. When we pass to more bulky organisms consisting of large numbers of protoplasts closely compacted together, the problem of relation to environment and of food transport become felt; the larger the organism usually the greater are these problems. A point is soon reached at which there is a gain by a differentiation in the work of different protoplasts, some for absorption, some for conduction, some for the light relation, some for reproduction, and so on. There is also a gain in splitting the form of the plant body up into parts so that a larger surface is exposed to environment with an economy in the amount of building material required. In this differentiation of the plant body into parts, there are two general problems to be solved, and the plant to be successful in its struggle for existence must control its development in such a way as to preserve the balance between them. (1) A ready display of a large surface to environment for the purpose of acquiring food and the disposition of waste. (2) The protection of the plant from injuries incident to an austere environment.

It is evident with the great variety of conditions met with in different parts of the same locality or region, and in different parts of the globe, that the plant has had very complex problems to meet and in the solution of them it has developed into a great variety of forms. It is also likely that different plants would in many cases meet these difficulties in different ways, sometimes with equal success, at other times with varied success. Just as different persons, given some one piece of work to do, are likely to employ different methods and reach results that are varied as to their value. While we cannot attribute consciousness or choice to plants in the sense in which we understand these qualities in higher animals, still there is something in their “constitution” or “character” whereby they respond in a different manner to the same influences of environment. This is, perhaps, imperceptible to us in the different individuals of the same species, but it is more marked in different species. Because of our ignorance of this occult power in the plant, we often speak of it as an “inherent” quality.

Perhaps the most striking examples one might use to illustrate the different line of organization among plants in two regions where the environment is very different are to be found in the adaptation of the cactus or the yucca to desert regions, and the oak or the cucurbits to the land conditions of our climate. The cactus with stem and leaf function combined in a massive trunk, or the yucca with bulky leaves expose little surface in comparison to the mass of substance, to the dry air. They have tissue for water storage and through their thick epidermis dole it out slowly since there is but little water to obtain from dry soil.

The cucurbits and the oak in their foliage leaves expose a very large surface in proportion to the mass of their substance, to an atmosphere not so severely dry as that of the desert, while the roots are able to obtain an abundant supply of water from the moist soil. The cactus and the yucca have differentiated their parts in a very different way from the oak or the cucurbits, in order to adapt themselves to the peculiar conditions of the environment.

When we say that certain plants have the power to adapt themselves to certain conditions of environment, we do not mean to say that if the cucurbits were transferred to the desert they would take on the form of the cactus or the yucca. They could do neither. They would perish, since the change would be too great for their organization. Nor do we mean, that, if the cactus or yucca were transferred from the desert to our climate, they would change into forms with thin foliage leaves. They could not. The fact is that they are enabled to live in our climate when we give them some care, but they show no signs of assuming characters like those of our vegetation. What we do mean is, that where the change is not too great nor too sudden, some of the plants become slightly modified. This would indicate that the process of organization and change of form is a very slow one, and is therefore a question of time—ages it may be—in which change in environment and adaptation in form and structure have gone on slowly hand in hand.

=690. Members of the plant body.=—The different parts into which the plant body has become differentiated are from one point of view, spoken of as members. It is evident that the simplest forms of life spoken of above do not have members. It is only when differentiation has reached the stage in which certain more or less prominent parts perform certain functions for the plant that members are recognized. In the algæ and fungi there is no differentiation into stem and leaf, though there is an approach to it in some of the higher forms. Where this simple plant body is flattened, as in the sea-wrack, or ulva, it is a _frond_. The Latin word for frond is _thallus_, and this name is applied to the plant body of all the lower plants, the algæ and fungi. The algæ and fungi together are sometimes called _thallophytes_, or _thallus plants_. The word thallus is also sometimes applied to the flattened body of the liverworts. In the foliose liverworts and mosses there is an axis with leaf-like expansions. These are believed by some to represent true stems and leaves; by others to represent a flattened thallus in which the margins are deeply and regularly divided, or in which the expansion has only taken place at regular intervals.

In the higher plants there is usually great differentiation of the plant body, though in many forms, as in the duckweeds, it is in the form of a frond. While there is a great variety in the form and function of the members of the plant body, they are all reducible to a few fundamental members. Some reduce these forms to three, the _root_, _stem_, _leaf_; while others to two, the _root_, and _shoot_, which is perhaps the best primary subdivision, and the shoot is then divided into stem and leaf, the leaf being a lateral outgrowth of the stem, and can be indicated by the following diagram:

{Stem. {Shoot····{ Plant body····{ {Leaf. {Root.

KINDS OF SHOOTS.

=691.= Since it is desirable to consider the shoot in its relation to environment, for convenience in discussion we may group shoots into four prominent kinds: (1) _Foliage shoots_; (2) _Shoots without foliage leaves_; (3) _Floral shoots_; (4) _Winter conditions of shoots and buds._ Topic (4) will be treated in Chapter XXXIX, section IV.

=692. (1st) Foliage shoots.=—Foliage shoots are either aerial, when their relation is to both light and air; or they are aquatic, when their relation is to both light and water. They bear green leaves, and whether in the air or water we see that light is one of the necessary relations for all. Naturally there are several ways in which a shoot may display its leaves to the light and air or water. Because of the great variety of conditions on the face of the earth and the multitudinous kinds of plants, there is the greatest diversity presented in the method of meeting these conditions. There is to be considered the problem of support to the shoot in the air, or in the water. The methods for solving this problem are fundamentally different in each case, because of the difference in the density of air and water, the latter being able to buoy up the plant to a great degree, particularly when the shoot is provided with air in its intercellular spaces or air cavities. In the solution of the problem in the relation of the shoot to aerial environment, stem and leaf have in most cases coöperated;[38] but in view of the great variety of stems and their modifications, as well as of leaves, it will be convenient to discuss them in separate chapters.

=693. (2d) Shoots without foliage leaves.=—These are subterranean or aerial. Nearly all subterranean shoots have also aerial shoots, the latter being for the display of foliage leaves (foliage-shoots), and also for the display of flowers (flower-shoots). The subterranean kinds bear scale leaves, i.e., the leaves not having a light relation are reduced in size, being small, and they lack chlorophyll. Examples are found in Solomon’s seal, mandrake (fig. 413_a_), etc. Here the scale leaves are on the bud at the end of the underground stem from which the foliage shoot arises. Aerial shoots which lack foliage leaves are the dodder, Indian-pipe-plant, beech drops, etc. These plants are saprophytes or parasites (see Chapter IX). Deriving their carbohydrate food from other living plants, or from humus, they do not need green leaves. The leaves have, therefore, probably been reduced in size to mere scales, and accompanying this there has been a loss of the chlorophyll. Other interesting examples of aerial shoots without foliage leaves are the cacti where the stem has assumed the leaf function and the leaves have become reduced to mere spines. The various modifications which shoots have undergone accompanying a change in their leaf relation will be discussed under stems in Chapter XXXIX.

=694. (3d) Floral shoots.=—The floral shoot is the part of the plant bearing the flower. As interpreted here it may consist of but a single flower with its stalk, as in Trillium, mandrake, etc., or of the clusters of flowers on special parts of the stem, termed flower clusters, as the _catkin_, _raceme_, _spike_, _umbel_, _head_, etc. In the floral shoot as thus interpreted there are several peculiarities to observe which distinguish it from the foliage shoot and adapt it to its life relations.

The floral shoot in many respects is comparable to the foliage shoot, as seen from the following peculiarities:

(1) It usually possesses, beside the flowers, small green leaves which are in fact foliage though they are very much reduced in size, because the function of the shoot as a foliage shoot is subordinated to the function of the floral shoot. These small leaves on the floral shoot are termed _bracts_.

(2) It may be (_a_) unbranched, when it would consist of a single flower, or (_b_) branched, when there would be several to many flowers in the flower cluster.

(3) The flower bud has the same origin on the shoot as the leaf bud; it is either terminal or axillary, or both.

(4) The members of the flower belong to the leaf series, i.e., they are leaves, but usually different in color from foliage leaves, because of the different life relation which they have to perform. Evidence of this is seen in the transition of sepals, petals, stamens, or pistils, to foliage leaves in many flowers, as in the pond lily, the abnormal forms of trillium, and many monstrosities in other flowers (see Chapter XXXIV).

(5) The position of the members of the flower on its axis, though usually more crowded, in many cases follows the same plan as the leaves on the stem.

The various kinds of floral shoots or flower clusters will be discussed in Chapter XLII, on the Floral Shoot.

II. Organization of Plant Tissues.

=695.= A tissue is a group of cells of the same kind having a similar position and function. In large and bulky plants different kinds of tissue are necessary, not only because the work of the plant can be more economically performed by a division of labor, but also cells in the interior of the mass or at a distance from the source of the food could not be supplied with food and air unless there were specialized channels for conducting food and specialized tissue for support of the large plant body. In these two ways most of the higher plants differ from the simple ones. The tissues for conduction are sometimes called collectively the _mestome_, while tissues for mechanical support are called _stereome_. Division of labor has gone further also so that there are special tissues for absorption, assimilation, perception, reproduction, and the like. The tissues of plants are usually grouped into three systems: (1) The Fundamental System, (2) The Fibrovascular System, (3) The Epidermal System. Some of the principal tissues are as follows:

1. THE FUNDAMENTAL SYSTEM.

=696. Parenchyma.=—Tissue composed of thin-walled cells which in the normal state are living. Parenchyma forms the loose and spongy tissue in leaves, as well as the palisade tissue (see Chapter IV); the soft tissue in the cortex of root and stem (Fig. 414); as well as that of the pith, of the pith-rays or medullary rays of the stem; and is mixed in with the other elements of the vascular bundle where it is spoken of as wood parenchyma and bast parenchyma; and it also includes the undifferentiated tissue (meristem) in the growing tips of roots and shoots; also the “intrafascicular” cambium (i.e., between the bundles, some also include the cambium within the bundle).

=697. Collenchyma.=—This is a strengthening tissue often found in the cortex of certain shoots. It also is composed of living cells. The cells are thickened at the angles, as in the tomato and many other herbs (fig. 414).

=698. Sclerenchyma, or stone-tissue.=—This is also a strengthening tissue and consists of cells which do not taper at the ends and the walls are evenly thickened, sometimes so thick that the inside (lumen) of the cell has nearly disappeared. Usually such cells contain no living contents at maturity. Sclerenchyma is very common in the hard parts of nuts, and underneath the epidermis of stems and leaves of many plants, as in the underground stems of the bracken fern, the leaves of pines (fig. 415), etc.

=699. Cork.=—In many cases there is a development of “cork” tissue underneath the epidermis. Cork tissue is developed by repeated division of parenchyma cells in such a way that rows of parallel cells are formed toward the outside. These are in distinct layers, soon lose their protoplasm and die; there are no intercellular spaces and the cells are usually of regular shape and fit close to each other. In some plants the cell walls are thin (cork oak), while in others they are thickened (beech). The tissue giving rise to cork is called “cork cambium,” or phellogen, and may occur in other parts of the plant. For example, where plants are wounded the living exposed parenchyma cells often change to cork cambium and develop a protective layer of cork. The walls of cork cells contain a substance termed _suberin_, which renders them nearly waterproof.

=700. Lenticels.=—These are developed quite abundantly underneath stomates on the twigs of birch, cherry, beech, elder, etc. The phellogen underneath the stoma develops a cushion of cork which presses outward in the form of an elevation at the summit of which is the stoma (fig. 416). The lenticels can easily be seen.

2. THE FIBROVASCULAR SYSTEM.

=701. Fibrous tissue.=[39]—This consists of thick-walled cells, usually without living contents which are elongated and taper at the ends so that the cells, or fibers, overlap. It is common as one of the elements of the vascular bundles, as wood fibers and bast fibers.

=702. Vascular tissue, or tracheary tissue.=—This consists of the vessels or ducts, and tracheides, which are so characteristic of the vascular bundle (see Chapter V) and forms a conducting tissue for the flow of water. The vascular tissue contains spiral, annular, pitted, and scalariform vessels and tracheides according to the marking on the walls (figs. 58, 59). These are all without protoplasmic contents when mature. There are also thin-walled living cells intermingled called wood parenchyma. In the conifers (pines, etc.) the tracheary tissue is devoid of true vessels except a few spiral vessels in the young stage, while it is characterized by tracheides with peculiar markings. These marks on the tracheides are due to the “bordered” pits appearing as two concentric rings one within the other. These can be easily seen in a longitudinal section of wood of conifers.

=703. Sieve tissue.=—This consists of elongated tubular cells connected at the ends, the cross walls being perforated at the ends. These are in the phloem part of the bundle, and serve to conduct downwards the dissolved substances elaborated in the leaves.

=704. Fascicular cambium.=—This is the living, cell-producing tissue in the vascular bundle, which in the open bundle adds to the phloem on one side and the xylem on the other.

3. THE EPIDERMAL SYSTEM.

=705.= To the epidermal system belong the epidermis and the various outgrowths of its cells in the form of hairs, or _trichomes_, as well as the guard cells of the stomates, and probably some of the reproductive organs.

=706. The epidermis.=—The epidermis proper consists of a single layer of external cells originating from the outer layer of parenchyma cells at the growing apex of the stem or root. These cells undergo various modifications of form. In many cases they lose their protoplasmic contents. In many cases the outer wall becomes thickened, especially in plants growing in dry situations or where they are exposed to drying conditions. The epidermal cells generally become considerably flattened, and are usually covered with a more or less well developed waterproof cuticle, a continuous layer over the epidermis. In many plants the cuticle is covered with a waxy exudation in the form of a thin layer, or of rounded grains, or slender rods, or grains and needles in several layers. These waxy coverings are sometimes spoken of as “bloom” on leaves and fruit.

=707. Trichomes.=—Trichome is a general term including various hair-like outgrowths from the epidermis, as well as scales, prickles, etc. These include root hairs, rhizoids, simple or branched hairs, glandular hairs, glandular scales, etc. Glandular hairs are found on many plants, as tomato, verbena, primula, etc.; glandular scales on the hop; simple-celled hairs on the evening primrose, cabbage, etc.; many-celled hairs on the primrose, pumpkin; branched hairs on the shepherd’s-purse, mullein, etc., stellate hairs on some oak leaves.

For stomates see Chapter IV.

4. ORIGIN OF THE TISSUES.

=708. Meristem tissue.=—The various tissues consisting of cells of dissimilar form are derived from young growing tissue known as _meristem_. Meristem tissue consists of cells nearly alike in form, with thin cell walls and rich in protoplasm. It is situated at the growing regions of the plants. In the higher plants these regions in general are three in number, the stem and root apex, and the cambium cylinder beneath the cortex. Tissues produced from the stem and root apex are called _primary_, those from the cambium _secondary_. In most cases the main bulk of the plant is secondary tissue, while in the corn plant it is all primary.

=709. Origin of stem tissues.=—Just back of the apical meristem in a longitudinal section of a growing point it can be seen that the cells are undergoing a change in form, and here are organized three formative regions. The outer layer of cells is called _dermatogen_ (skin producer), because later it becomes the epidermis. The central group of elongating cells is the _plerome_ (to fill). This later develops the _central cylinder_, or _stele_, as it is called (fig. 417). Surrounding the plerome and filling the space between it and the dermatogen is the third formative tissue called the _periblem_, which later forms the cortex (bark or rind), and consists of parenchyma, collenchyma, sclerenchyma, or cork, etc., as the case may be. It should be understood that all these different forms and kinds of cells have been derived from meristem by gradual change. In the mature stems, therefore, there are three distinct regions, the central cylinder or stele, the cortex, and the epidermis.

=710. Central cylinder or stele.=—As the central cylinder is organized from the plerome it becomes differentiated into the vascular bundles, the pith, the pith-rays (medullary rays) which radiate from the pith in the center between the bundles out to the cortex, and the pericycle, a layer of cells lying between the central cylinder and the cortex. The bundles then are farther organized into the xylem and phloem portions with their different elements, and the fascicular cambium (meristem) separating the xylem and phloem, as described in Chapter V. Such a bundle, where the xylem and phloem portions are separated by the cambium is called an open bundle (as in fig. 58). Where the phloem and xylem lie side by side in the same radius the bundle is a _collateral_ one. Dicotyledons and conifers are characterized by open collateral bundles. This is why trees and many other perennial plants continue to grow in diameter each year. The cambium in the open bundle forms new tissue each spring and summer, thus adding to the phloem on the outside and the xylem on the inside. In the spring and early summer the large vessels in the xylem predominate, while in late summer wood fibers and small vessels predominate and this part of the wood is firmer. Since the vascular bundles in the stem form a circle in the cylinder, this difference in the size of the spring and late summer wood produces the “annual” rings, so evident in the cross-section of a tree trunk. Branches originate at the surface involving epidermis, cortex, and the bundles.

In monocotyledonous plants (corn, palm, etc.) the bundles are not regularly arranged to form a hollow cylinder, but are irregularly situated through the stele. There is no meristem, or cambium, left between the xylem and phloem portions of the bundle and the bundle is thus _closed_ (as in fig. 60), since it all passes over into permanent tissue. In most monocotyledons there is, therefore, practically no annual increase in diameter of the stem.

=711. Ferns.=—In the ferns and most of the Pteridophytes an apical meristem tissue is wanting, its place being taken by a single apical cell from the several sides of which cells are successively cut off, though Isoetes and many species of Lycopodium have an apical meristem group. In most of the Pteridophytes also the bundles are _concentric_ instead of collateral. Fig. 418 represents one of the bundles from the stem of the polypody fern. The xylem is in the center, this surrounded by the phloem, the phloem by the phloem sheath, and this in turn by the endodermis, giving a concentric arrangement of the component tissues. A cross-section of the stem (fig. 419) shows two large areas of sclerenchyma, which gives the chief mechanical support, the bundles being comparatively weak.

=712. Origin of root tissues.=—A similar apical meristem exists in roots, but there is in addition a fourth region of formative tissue in front of the meristem called _calyptrogen_ (fig. 420). This gives rise to the “root cap” which serves to protect the meristem. The vascular cylinder in roots is very different from that of the stem. There is a solid central cylinder in which the groups of xylem radiate from the center and groups of phloem alternate with them but do not extend so near the center (fig. 421). As the root ages, changes take place which obscure this arrangement more or less. Branches of the roots arise from the central cylinder. In fern roots the apical meristem is replaced by a single four-sided (tetrahedral) apical cell, the root cap being cut off by successive divisions of the outer face, while the primary root tissues are derived from the three lateral faces.

=Function of the root cap.=—The root cap serves an important function in protecting the delicate meristem or cambium at the tip of the root. These cells are, of course, very thin-walled, and while there is not so much danger that they would be injured from dryness, since the soil is usually moist enough to prevent evaporation, they would be liable to injury from friction with the rough particles of soil. No similar cap is developed on the end of the stem, but the meristem here is protected by the overlapping bud-scales. One of the most striking illustrations of a root cap may be seen in the case of the Pandanus, or screw-pine, often grown in conservatories (see fig. 447). On the prop roots which have not yet reached the ground the root caps can readily be seen, since they are so large that they fit over the end of the root like a thimble on the finger.

=713. Descriptive Classification of Tissues.=

{ Epidermis. { { { Simple hairs. { { Many-celled hairs. { { Branched hairs, often stellate. Epidermal { Trichomes. { Clustered, tufted hairs. System. ····{ { Glandular hairs. { { Root hairs. { { Prickles. { { Guard cells of stomates. { Spiral vessels. { Pitted vessels. { Scalariform vessels. { Xylem (wood). { Annular vessels. { { Tracheides. { { Wood fibers. { { Wood parenchyma. Fibrovascular { System. ····{ Cambium (fascicular). { { { Sieve tubes. { Phloem (bast). { Bast fibers. { Companion cells. { Bast parenchyma. { Cork. { Collenchyma. { Cortex.····{ Parenchyma. { { Fibers. { { Milk tissue. { { Pith-ray.··{ Parenchyma. { { Intrafascicular { Stem and root. { cambium. { { Pith.······{ Parenchyma. { { { Sclerenchyma. { { Fundamental { { Bundle-sheath. System. ····{ { { { Endodermis. { { Leaves. { Palisade tissue. { { Spongy parenchyma. { { Reproductive Organs (mainly fundamental).

=714. Physiological Classification of Tissues.=

_Formative Tissue._

Thin-walled cells composing the meristem, capable of division and from which other tissues are formed.

_Protective Tissue._

_Tegumentary System._—Epidermis, periderm, bark protecting the plant from external contact.

_Mechanical System._—Bast tissue, bast-like tissue, collenchyma, sclerenchyma, afford protection against harmful bending, pulling, etc.

_Nutritive Tissues._

_Absorptive System._—Root hairs and cells, rhizoids, aerial root tissue, absorptive leaf glands, absorptive organs in seeds, haustoria of parasites, etc.

_Assimilatory System._—Assimilating cells in leaf and stem.

_Conductive System._—Sieve tissue, tracheary tissue, milk tissue, conducting parenchyma, etc.

_Food-storing System._—Water reservoir, water tissue, slime tissue, fleshy roots and stems, endosperm and cotyledons, etc.

_Aerating System._—Air spaces and tubes, special air tissue, air-seeking roots, stomates, lenticels, etc.

_Secretory and Excretory System._—Water glands, digestive glands, resin glands, nectaries, tannin, pitch and oil receptacles, etc.

_Apparatus and Tissues for Special Duties._

Holdfasts.

Tissues of movement, parachute hairs, floating tissue, hygroscopic tissue, living tissue.

For perceiving stimuli.

For conducting stimuli, etc.

FOOTNOTES:

[37] =Suggestions to the teacher.=—In the study of the flowering plants in the secondary school and in elementary courses three general topics are suggested. 1st, the study of the form and members of the plant and their arrangement, as in Chapters XXXVIII-XLV. 2d, the study of a few plants representative of the more important families, in order that the members of the plant, as studied under the first topic, may be seen in correlation with the plant as a whole in a number of different types. 3d, the study of plants in their relation to environment, as in Chapter XLVI. The first and second topics can be conducted consecutively in the classroom and laboratory. The third topic can be studied at opportune times during the progress of topics 1 and 2. For example, while studying topic 1 excursions can be made to study winter conditions of buds, shoots, etc., if in winter period, or the relations of leaves, etc., to environment, if in the growing period. While studying topic 2 excursions can be made to study flower relations, and also vegetation relations to environment (see Chapters XLVI-LVII of the author’s “College Text-book of Botany”). It is believed that a study of these three general topics is of much more value to the beginning student than the ordinary plant analysis and determination of species.

[38] It is interesting to note that in some foliage shoots the stem is entirely subterranean. See discussion of the bracken fern and sensitive fern in Chapter XXXIX.

[39] Some fibers occur also very frequently in the Fundamental System, forming bundle-sheaths, or strands of mechanical tissue in the cortex.