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Originally appearing in Volume V23, Page 126 of the 1911 Encyclopedia Britannica.
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REPRODUCTION OF PLANTS The various modes in which plants reproduce their species may be conveniently classified into two groups, namely, vegetative propagation and true reproduction, the distinction between them being roughly this, that whereas in the former the production of the new individual may be effected by the most various parts of the body, in the latter it is always effected by means of a specialised reproductive cell. I. Vegetative Propagation. The simplest case of vegetative multiplication is afforded by unicellular plants. When the cell which constitutes the body of the plant has attained its limit of size it gives rise to two either by division or gemmation; the two cells then grow, and at the same time become separated from each other, so that eventually two new distinct individuals are produced, each of which precisely resembles the original organism. A good example of this is to be found in the germination of the yeast plant. ' This mode of multiplication is simply the result of the ordinary processes of growth. All plant-cells grow and divide at some time or other of their life; but whereas in multicellular plant the products of division remain coherent, and add to the number of the cells of which the plant consists, in a unicellular plant they separate and constitute new individuals. In more highly organized plants vegetative propagation may be effected by the separation of the different parts of the body from each other, each such part developing the missing members and thus constituting a new individual. This takes place spontaneously in rhizomatous plants, in which the main stem gradually dies away from behind forwards; the lateral branches thus become isolated and constitute new individuals. The remarkable regenerative capacity of plant-members is largely made use of for the artificial propagation of plants. A branch removed from a parent-plant will, under appropriate conditions, develop roots, and so constitute a new plant; this is the theory of propagation by " cuttings." A portion of a root will similarly develop one or more shoots, and thus give rise to a new plant. An isolated leaf will, in many cases, produce a shoot and a root, that is, a new plant; it is in this way that new begonias, for instance, are propagated.. The production of plants from leaves occurs also in nature, as, for instance, in certain so-called " viviparous plants, of which Bryophyllum calycinum (Crassulaceae) and many ferns [Nephrodium (Lastraea) Filix-mas, Asplenium (Athyrium) •Filix foemina and other species of Asplenium] are examples. But it is in the mosses, of all plants, that the capacity for vegetative propagation is most widely diffused. Any part of a moss, whether it be the stem, the leaves, the rhizoids, or the sporogonium, is capable, under appropriate conditions, of giving rise to _filamentous protonema, on which new moss-plants are then developed as lateral buds. In a large number of plants provision is made for vegetative propagation by the development of more or less highly specialized organs. In lichens, for instance, there are the soredia, which are minute buds of the thallus containing both algal and fungal elements; these are set free on the surface in large numbers, and each grows into a thallus. In the Characeae there are the bulbils or " starch-stars " of Chara stelligera, which are under-ground nodes, and the branches with naked base and. the pro-embryonic branches found by Pringsheim on old nodes of Chara fragilis. In the mosses small tuberous bulbils frequently occur on the rhizoids, and in many instances (Bryum annotinum, Aulacomnion androgynum, Tetraphis pellucida, &c.) stalked fusiform or lenticular multicellular bodies containing chlorophyll, termed gemmae, are produced on the shoots, either in the axils of the leaves or in special receptacles at the summit of the stem. Gemmae of this kind are produced in vast numbers in Marchantia and Lunularia among the liverworts. Similar gernmae are also produced by the prothallia of ferns. In some ferns (e.g. Nephrolepis tuberosa and undulata) the buds borne on the leaves or in their axils become swollen and filled with nutritive materials, constituting bulbils which fall off and give rise to new plants. This conversion of buds into bulbils, which subserve vegetative multiplication, occurs also occasionally among Phanerogams, as for instance in Lilium bulbiferum, species of Poet, Polygonum viviparum, &c. But many other adaptations of the same kind occur among Phanerogams. Bulbous plants, for instance, produce each year at least one bulb or corm from which a new plant is produced in the succeeding year. In the potato, tubers are developed from subterranean shoots, each of which in thefollowing year gives rise to a new individual. In the dahlia, Thladiantha dubia, &c., tuberous swellings are found on the roots, from each of which a new individual may spring. II. True Reproduction. This is effected by cells formed by the proper reproductive organs. These cells are of two principal kinds. There are, first, those cells each of which is capable of developing by itself into a new organism: these are the asexual reproductive cells, known generally as spores. Secondly, there are the cells which are incapable of independent germination; it is not until these cells have fused together in pairs that a new organism can be developed: these are the sexual reproductive cells or gametes. In some exceptional cases the normal mode of reproduction, sexual or asexual, does not take place: instead, the new organism is developed vegetatively from the parent. When sexual reproduction is suppressed the case is one of apogamy; when asexual reproduction by spores is suppressed the case is one of apospory. (Apogamy and apospory are discussed below in the section on Abnormalities of Reproduction.) Asexual Reproduction.—Reproduction by means of some kind of spore (using the term in its widest sense, so as to include all asexually produced reproductive cells) is common to nearly all families of plants; it is wanting in certain Algae (Conjugatae, Fucaceae, Characeae), and in certain fungi (e.g. some Peronosporeae). The structure of a spore is essentially this: it consists of a nucleated mass of protoplasm, enclosing starch or oil as re-serve nutritive material, usually invested by a cell-wall. In those cases in which the spore is capable of germinating immediately on its development the cell-wall is a single delicate membrane consisting of cellulose; but in those cases in which the spore may or must pass through a period of quiescence before germination the wall becomes thickened and may consist of two layers, an inner, the endospore, which is delicate and consists of cellulose, and an outer, the exospore, which is thick and rigid, frequently darkly coloured and beset externally with spines or bosses, and which consists of cutin. In some few cases among the fungi, multicellular or septate spores are produced; these approximate somewhat to the gemmae mentioned above as highly specialized organs for vegetative propagation. In some cases, particularly among the algae, and also in some fungi (Peronosporeae, Saprolegnieae, Chytridiaceae, and the Myxomycetes), spores are produced which are usually destitute of any cell-wall, and are further peculiar in that they are motile, and are therefore termed zoospores; they move sometimes in an amoeboid manner by the protrusion of pseudopodia, but more frequently they are provided with one, two, or many delicate vibratile protoplasmic filaments, termed cilia, by the lashing of which the spore is propelled through the water. The zoospore eventually comes to rest, withdraws its cilia, surrounds itself with a cell-wall, and then germinates. In the simplest case a single spore is developed from the cell of the unicellular plant, the protoplasm of which surrounds itself with the characteristic thick wall. This occurs only in plants of low organization such as the Schizophyta. In other cases the contents of the cell undergo division, each portion of the protoplasm constituting a spore. Examples of this are afforded, among unicellular plants, by yeast and the Protococcaceae; and in multicellular plants by the Pandorineae, Confervaceae, Ulvaceae, &c., where any cell of the body may produce spores. In such cases the spore-producing cell may be regarded as a rudimentary reproductive organ of the nature of a sporangium. In more highly organized plants special organs are differentiated for the production of spores. In the majority of cases the special organ is a sporangium, that is, a capsule in the interior of which the spores are developed; but in many fungi the spores are formed by abstriction from an organ termed a sporophore. In the Thallophyta the sporangium is commonly a single cell. In the Bryophyta it is a multicellular capsule. In the Pteridophyta the sporangium is multicellular, but simple in structure. and this is true also of the Phanerogams. It is important to note that in all the Bryophyta and in some of the Pteridophyta (most of the Filicinae, all existing Equisetinae, and the Lycopodiaceae and Psilotaceae) there is but one kind of sporangium and spore, the plants being homosporous or isosporous, whereas the rest of the Pteridophyta (Hydropterideae, Selaginellaceae) and the Phanerogams are heterosporous, having sporangia of two kinds; some produce one or a few large spores (megaspores), and are hence termed mega-sporangia, while others give rise to a larger number of small spores (microspores) and are hence termed microsporangia. In the Phanerogams the two kinds of sporangia have received special names: the megasporangium, which produces as a rule only one mature spore (embryo-sac), is termed the ovule; the microsporangium, which produces a large number of micro-spores (pollen-grains), is termed the pollen-sac. The development of spores, except in the simpler Thallophyta, is more or less restricted to definite parts of the body. Thus in the Red Algae (Florideae) there are the organs known as stichidia, nemathecia. In the fungi the number and variety of such organs is very great; they may be described generally as simple and compound sporophores: but for a description the article FUNGI should be consulted. In the higher plants the organs are less various. In the Bryophyta the production of spores is restricted to the sporogonium. In the vascular plants (Pteridophyta, Phanerogams) the development of sporangia, speaking generally, is confined to the leaves. In most ferns the sporangiferous leaves (sporophylls) do not differ in appearance from the foliage leaves; but in other Pteridophyta (Equisetaceae, Marsiliaceae, some species of Lycopodium and Selaginella) they present considerable adaptation, and notably in the Phanerogams. In the Phanerogams the specialization is so great that the sporophylls have received special names; those which bear the microsporangia (pollen-sacs) are termed the stamens, and those which bear the megasporangia (ovules) are termed the carpels. The sporophylls are usually aggregated together on a short stem, forming a shoot that constitutes a flower. Many terms are employed to indicate the nature of the various kinds of spores, especially among the fungi, but the endless varieties of asexual (and asexually produced) reproductive cells may be grouped under two heads—0) Gonidia, (2) Spores proper. The distinction between these two kinds of asexual reproductive cells is as follows. The gonidium is a reproductive cell that gives rise, on germination, to an organism resembling the parent. For instance, among the algae, the " zoospore " of Vaucheria develops into a ,Vaucheria-plant. There is thus a close connexion between vegetative multiplication and multiplication by means of gonida. The production of gonida is entirely limited to the Thallophyta, and is especially marked in the fungi, though the nature of all the many kinds of reproductive cells formed in this group has not yet been fully investigated. It is, however, wanting in certain algae (Conjugatae, Fucaceae, Characeae) and fungi (some Peronosporeae and Ascomycetes). The spore proper is a reproductive cell that as a rule gives rise, on germination, to an organism unlike that which produced it. For instance, the spore of a fern when it germinates gives rise, not to a fern-plant, but to a prothallium. The apparent exceptions to this rule occur only among the Thallophyta, and are explained below in the section on Life-history. The true spore is developed, usually in a sporangium, after a process of division which presents certain features that call for special notice. Observation of the process of division of the nucleus (karyokinesis) in plants generally has shown (for details see CYTOLOGY) that the limn-reticulum of the resting nucleus breaks up into a definite number of segments, the chromosomes, each of which bears a series of minute bodies, the chromatin-disks or chromomeres, consisting largely of a substance termed chromatin. In the ordinary homotype divisions of the nuclei the characteristic number of chromosomes is always observable: but when the spore-mother-cells are being formed the number of chromosomesis reduced to one-half. This, if the number of chromosomes of the parent plant be expressed as 2X the number in the spore will be x. To take a concrete case: it has been observed by Guignard and others that in the early divisions taking place in the developing anther and ovule of the lily the number of chromosomes is 24; whereas in the later divisions which give rise to the pollen-mother-cells in the one case and to the mother-cell of the embryo-sac in the other, the number of chromosomes is only 12. Thus the development of a spore (as distinguished from a gonidium) is always preceded by a reducing- or heterotypedivision, a process now more generally termed meiosis (Farmer). The reduced number of chromosomes in the nucleus of the spore-mother-cell persists in the spore, and in all the cells of the organism to which the spore may give rise. (Meiosis is discussed below in the section on Sexual Reproduction.) It should be explained that cells, to which the name " spore " has also been applied, are formed as the result of a sexual act: such are zygospores, oospores, and some carpospores. But these cells differ from spores proper not only in their mode of origin but also in that their nuclei contain the full double number (2x) of chromosomes; hence they may be distinguished as diplospores. Sexual Reproduction.—Sexual reproduction involves the development of sexual organs (gametangia) and sexual cells (gametes). When the organism is unicellular, as in the lower Green Algae (e.g. Protococcaceae, Conjugatae), the cell becomes a sexual organ and its whole protoplasm gives rise to one or more sexual cells: in the higher forms certain parts of the body are specialized as sexual organs. In many of the lower plants the organs present no external distinction of sex (e.g. lower Green Algae: the Chytridiaceae, Mucorinae, and some Ascomycetes among the fungi): it is impossible to distinguish between the male and female organs, although it cannot be doubted that the essential physiological difference exists; consequently the organs are merely described as gametangia. The gap between these plants and those with differentiated sexual organs is, however, bridged over by intermediate forms, as explained in the article ALGAE. When the sexual organs are more or less obviously differentiated into male and female, they present considerable variety of form in different groups of plants, and accordingly bear different names. Thus the male organ is a pollinodium in most of the fungi, a spermogonium in others (certain Ascomycetes, Uredineae); in all other plants it is an antheridium. Similarly the female organ is an oogonium in various Thallophyta (Green and Brown Algae: Oomycetous Fungi); a procarp in the Red Algae; an archicarp in certain Ascom+ycetous Fungi and in the Uredineae; an archegonium in all the higher plants. It is generally the case that the protoplasm of the sexual organ is differentiated into one or more sexual cells. Thus, the gametangium usually gives rise to cells which, as they are externally similar, are termed isogametes or simply gametes. Certain forms of the male organ, the spermogonium and the antheridium, give rise to male cells which are termed spermatia when they are non-ciliate, spermatozoids when they are ciliated and free-swimming.. Again, the female organs termed oogonia and archegonia produce one or more female cells called oospheres. But there are important exceptions to this rule. Thus the protoplasm is not differentiated into cells in the gametangium of the Mucorinae; in the male organ (pollinodium), of fungi generally; and in the female organ (procarp) of the Red Algae and (archicarp) of the Ascomycetes and Uredineae. The immediate product of the fusion of cells, or of undifferentiated protoplasm, derived from sexual organs of opposite sex may be generally termed the zygote; but it is not always of the same kind. Thus when two isogametes, or the undifferentiated contents of two gametangia, fuse together, the process is designated conjugation, and the product is usually a single cell termed zygospore. When an oosphere fuses with a male cell, or with the undifferentiated contents of a male organ, the process is fertilization, and the product is a single cell termed oospore. When, finally, a female organ with undifferentiated contents receives a male cell, the process again is fertilization; here the product is not a single cell, but a fructification termed cystocarp (Red Algae), or ascocarp (Ascomycetes) or aecidium (Uredineae), containing many spores (car pospores). As a consequence of the diversity in the sexual organs and cells, in the details of the sexual act, and in the product of it, several modes of the sexual process have to be distinguished, which may be conveniently summarized as follows: I. Isogamy: the sexual process consists in the fusion of either two similar sexual cells (isogametes), or two similar sexual organs (gametangia): it is termed conjugation, and the product is a zygospore. Its varieties are: (a) Gametes ciliated and free-swimming (planogametes), set free into the water where they meet and fuse: lower Green Algae (Protococcaceae, Pandorineae, most Siphonaceae and Confervaceae) ; some Brown Algae (Phaeosporeae) : (b) Gametangia fuse in pairs, and a gamete is differentiated in each: the gametes of each pair fuse, but are not set free and are not ciliated (the Conjugate Green Algae): or, no gametes are differentiated, the undifferentiated con-tents of the gametangia fusing (Mucorinae among the Fungi). II. Oogamy: male and female organs distinct: the protoplasm of the female organ is differentiated into one or (rarely) more oospheres which usually remain enclosed in the female organ: the contents of the male organ are usually differentiated into one or more male cells: the process is fertilization, the product is an oos ore. A) The sexual organs are unicellular (or coenocytic as in certain Siphonaceous Green Algae and in the Oomycetous Fungi) ; the female organ is an oogonium. (a) The male organ is an antheridium giving rise to one or more free-swimming ciliated spermatozoids: (I) The oogonium contains a single oosphere which is fertilized in situ: higher Green Algae (Volvox, Vaucheria, Oedogonium, Coleochaete, Characeae) ; some Brown Algae (Tilopteris) ; among the Fungi, Monoblepharis, the only fungus known to have spermatozoids: (2) The oogonium produces a single oosphere which is extruded and is fertilized in the water: Dietyota and some Fucaceae (Brown Algae) : (3) The oogonium contains several oospheres which are fertilized in situ: Sphaeroplea (Siphonaceous Green Alga): (¢) The oogonium produces more than one oosphere (2–8) which are extruded and are fertilized in the water: certain Brown Algae (Pelvetia, Ascophyllum, Fucus): (9) The male organ is a pollinodium which applies itself closely to the oogonium: the amorphous male cell is not ciliated and is not set free: (I) The oogonium contains a single oosphere which is fertilized in situ: Peronosporaceae (Oomycetes): (2) The oogonium contains several oospheres; Saprolegniaceae: but it is debated whether or not fertilization actually takes place. (B) The male and female organs are (as a rule) multicellular; the male organ is an antheridium, the female an archegonium: the archegonium always contains a single oosphere which is fertilized in situ. (a) The male cell is a free-swimming ciliated spermatozoid: the antheridium produces more than one (usually very many) spermatozoids, each of which is developed in a single cell: all Bryophyta (mosses, &c.) and Pteridophyta (ferns, &c.): the only Phanerogams in which spermatozoids have been observed are the gymnospermous species Ginkgo biloba, Cycas revoluta, Zamia integrifolia. (') The male cell is amorphous and passes directly from the pollen-tube into the oosphere (siphonogamy) : all Phanerogams except the species just mentioned. It must be explained that in the angiospermous Phanerogams, the male and female organs are so reduced that each is represented by only a single cell: the male, by the generative cell, formed in the pollen-grain, which usually divides into two male cells: the female, by the oosphere. The gradual reduction can be traced through the Gymnosperms. Attention may here be drawn to the fact (see ANGIOSPERMS) that, in several cases, the second male cell has been seen to enter the embryo-sac from the pollen-tube, and its nucleus to fuse with the definitive nucleus (endosperm-nucleus) or with one of the polar nuclei. The significance of this remarkable observation is discussed in the section on the Physiology of Reproduction. (A) There are definite male cells (spermatia) : (a) The female organ is a procarp, consisting of an elongated, closed, receptive filament, the trichogyne, and of a basal fertile portion, the carpogonium: on fertilization the latter grows and gives rise directly or indirectly to a cystocarp: the spermatia are each formed in a unicellular antheridium and have no cell-wall at first: they fuse with the tip of the trichogyne : Red Algae (Rhodophyceae or Florideae): (~) The female organ (archicarp) resembles the preceding: in fertilization the fertile portion (ascogonium) develops into an ascocarp containing one or more asci (sporangia) each containing usually eight ascospores: the spermatia are formed by abstriction from the filaments (sterigmata) lining special receptacles, the spermogonia, which are the male organs: certain Ascomycetous Fungi (e.g. Laboulbeniaceae, some Lichen-Fungi, Polystigma). For the Uredineae, see Abnormalities of Reproduction, below). (B) There are no definite male cells: the more or less distinct male and female organs come into contact, and their undiffer- entiated contents fuse: the product is an ascocarp: (a) The male and female organs are obviously different: the female organ is an ascogonium, the male a pollinodium: e.g. Pyronema, Sphaerotheca (Ascomycetes) : (0) The male and female organs are quite similar: e.g. Eremascus, Dipodascus (Ascomycetes). It may be explained that carpogamy is the expression of sexual degeneration. In the cases last mentioned, when the sexual organs are quite similar, they have reverted to the condition of gametangia. Still further reduction is observable in other Ascomycetes in which one of the sexual organs, presumably the male, is either much reduced or is altogether wanting. Again in the rusts (Uredineae), there are spermatia, but they are functionless (see section on Abnormalities of Reproduction). In the highest Fungi, the Autobasidiomycetes, no sexual organs have been discovered. Details of the Sexual Act.—It has been already stated that the sexual act consists in the fusion of two masses of protoplasm, commonly cells, derived from two organs of opposite sex: but this is only the first stage in the process. The second stage is the fusion of the nuclei, which usually follows quickly upon the fusion of the cells; but nuclear fusion may be postponed so that the two sexual nuclei may be observed in the zygote, as " conjugate " nuclei, and even in the cells of the organism developed from the zygote (e.g. Uredineae). The result of nuclear fusion is that the nucleus of the zygote contains the double number of chromosomes—that is, if the number of chromosomes in each of the fusing sexual nuclei be x, the number in the nucleus of the zygote will be 2x. Moreover, this double number persists in all the cells of the organism developed from the zygote, until it is reduced to one-half by meiosis preceding either the development of the spores, or, less commonly, the development of the sexual cells. But there is yet a third stage, which consists in the temporary fusion of the chromosomes belonging to the two sexual nuclei. This always takes place as a preliminary to meiosis; it may be in the germinating zygote, or after many generations of cells have been formed from it. At the onset of meiosis the (2x) chromosomes are seen to be double, one of each pair having been derived from the male and the female cell respectively: the chromosomes of each pair then fuse so that their chromomeres unite along their length, constituting the pseudo-chromosomes. The paired chromosomes separate and eventually go to form the two daughter-nuclei, one to each, which thus have half (x) the original number of chromosomes. The daughter-nuclei at once divide homotypically, retaining the reduced (x) number of chromosomes to form the four nuclei of a tetrad of spores (more rarely, e.g. Fucus, of sexual cells). It will have been gathered from the foregoing sections that plants generally are capable of both sexual and asexual reproduction; and, further, that in different stages of their life-history they possess the diploid (2x) number of chromosomes in. their nuclei, or the haploid (x) number. It may be at once stated that, in all plants in which sexual reproduction and true meiotic spore-formation exist, these two modes of reproduction are restricted to distinct forms of the plant; the sexual form bears only the sexual organs and is haploid; the asexual form, only produces spores and is diploid. Hence all such plants are to this extent polymorphic—that is, the plant assumes these two forms in the course of its life-history. When, as in many Thallophyta, one or other of these forms can reproduce itself by means of gonidia, additional forms may be introduced into the life-history, which becomes the more complicated the more pronounced the polymorphism. The most straightforward life-histories are those presented by the Bryophyta and the Pteridophyta, where there are but the two forms, the sexual and the asexual. In the life-history of a moss, the plant itself bears only sexual organs: it is the sexual form, and is distinguished as the gametophyte. The zygote (oospore) formed in the sexual act develops into an organism, the sporogonium, which is entirely asexual, producing only spores: it is distinguished as the sporophyte. When these spores germinate, they give rise to moss-plants. Thus the two forms, the sexual and the asexual, regularly alternate with each other—that is, the life-history presents that simple form of polymorphism which is known as alternation of generations. Similarly, in the life-history of a fern, there is a regular alternation of a sporophyte, which is the fern-plant itself, with a gametophyte, which is the fern-prothallium. It is pointed out in the preceding section that, as the result of the sexual act, the nucleus of the zygote contains twice as many chromosomes as those of the fusing sexual cells. This 2X number of chromosomes persists throughout all the cell-generations derived from the zygote, that is, in the cells constituting the sporophyte, up to the time that it begins to produce spores, when meiosis takes place. Again, the cell-generations derived from the spore, that is, the cells constituting the gametophyte, all have the reduced x number of chromosomes in their nuclei up to the sexual act. Hence the sporophyte may also be designated the diplophyte and the gametophyte the haplophyte (Strasburger): in other words, the sporophyte is the pre-meiotic, the gametophyte the post-meiotic generation. Twice in its life-history the plant is represented by a single cell: by the spore and by the zygote. The turning-points in the life-history, the transitions from the one generation to the other, are (r) meiosis, (2) the sexual act. The course of the life-history in Phanerogams and in those Thallophyta which have been adequately investigated is essentially the same as that of the Bryophyta and of the Pteridophyta as described above, though it is less easy to trace on account of the peculiar relation of the two generations to each other in the Phanerogams and on account of various irregularities that present themselves in the Thallophyta. In the Phanerogams, as in the Pteridophyta, the preponderating generation is the sporophyte, the plant itself. Inasmuch as they are heterosporous, the gametophyte is represented by a male and a female organism or prothallium, both rudimentary. The male prothallium consists of the few cells formed by the germinating pollen-grain (microspore); and though it is quite independent, since the microspores are shed, it grows parasitically in the tissues upon which the microspore has been deposited in pollination. The female prothallium may consist of many cells with well-developed archegonia, as in the Gymnosperms, or of only a few cells with the female organ reduced to the oosphere, as in the Angiosperms. In either case it is the product of the germination of a megaspore (embryo-sac) which is not shed from its sporangium (ovule): hence it never becomes an independent plant, and was long regarded as merely a part of the sporophyte until its true nature was ascertained, chiefly by the researches of Hofmeister, who first explained the alternation of generations in plants. This intimate and persistent connexion between the two generations affords the explanation of the characteristic features of the Phanerogams, the seed and the flower. The ovule containing the embryo-sac, which eventually contains the embryo, persists as the seed—a structure that is distinctive of Phanerogams, which have, in fact, on this account been also termed Spermatophyta. With regard to the flower, it has been already mentioned that it is, like the cone of an Equisetum or a Lyco-podium, a shoot adapted to the production of spores. But it is something more than this: for whereas in Equisetum or Lycopodium the function of the cone comes to an end when the spores are shed, the flower of the Phanerogam has still various functions to perform after the maturation of the spores. It is the seat of the process of pollination—that is, the bringing of the pollen-grain by one of various agencies into such a position that a part (the pollen-tube) of the male prothallium developed from it may reach and fertilize the oosphere in the embryo-sac. Thus the flower of Phanerogams is a reproductive shoot adapted not only for spore-production, but also for pollination, for fertilization, and for the consequences of fertilization, the production of seed and fruit. However, in spite of these complications, it is possible to determine accurately the limits of the two generations by the observation of the nuclei. The meiosis preceding the formation of the spores marks the beginning of the (haploid) gametophyte, male and female; and the sexual act marks that of the (diploid) sporophyte. The difficult task of elucidating the life-histories of the Thallophyta has been successfully performed in certain cases by the application of the method of chromosome-counting, with the result that alternation of generations has been found to be of general occurrence. To begin with the Algae. In the Dictyotaceae (Brown Algae) there are two very similar forms in the life-history, the one bearing asexual reproductive organs (tetrasporangia), the other bearing sexual organs (oogonia ' and antheridia). It has 'been shown (Lloyd Williams) that the former is undoubtedly the sporophyte and the latter the gametophyte, since the nuclei of the former contain 32 chromosomes, and those of the latter 16. Meiosis takes place in the mother-cell of the tetraspores, which, on germination, give rise to the sexual form. Quite a different life-history has been traced in Fucus, another Brown Alga. Here no spores are produced: there is but one form in the life-history, the Fucusplant, which bears sexual organs and has, on that account, been regarded as a gametophyte. The investigation of the nuclei has, however, shown (Farmer) that the Fucus-plant is actually diploid, that it is, in fact, a sporophyte; but since there is no spore-formation, meiosis immediately precedes the development of the sexual cells, which alone represent the gametophyte (see below, Apospory). Similarly, two types of life-history have been discovered in the Red Algae. In Polysiphonia violacea, a species in which the tetraspores and the sexual organs are borne by similar but distinct individuals, it has been ascertained (Yamanouchi) that, as in Dictyota, meiosis takes place in the mother-cell of the tetraspores, so that the nuclei of these spores, as also those of the sexual plants to which they give rise, contain 20 chromosomes: and further, that the nuclei of the carpospores (diplospores) produced in the cystocarp as the result of fertilization, contain 40 chromosomes, as do also those of the asexual plant to which the carpospores give rise. Hence the sporophyte is represented by the cystocarp and the resulting tetrasporangiate plants: the gametophyte, by the sexual plants. Though it is the rule in the Red Algae that the tetrasporangia and the sexual organs are borne on distinct individuals, yet cases are known in which both kinds of reproductive organs are borne upon the same plant; and to those the above conclusions obviously cannot apply. They have yet to be investigated. The second type of life-history has been traced in Nemalion. Here there is no tetrasporangiate form, consequently meiosis takes place at a different stage in the life-history. It has been observed (Wolfe) that the nuclei of the sexual plant contain 8 chromosomes; those of the gonimoblast-filaments of the developing cystocarp contain 16, whilst those of the carpospores contain 8: hence meiosis takes place in the carposporangia. Here the plant is the gametophyte; the sporophyte is only represented by the cystocarp. The carpospores here are true spores (haplospores). Among the Green Algae, Coleochaete is the only form that has been fully investigated (Allen). Here meiosis takes place in the germinating oospore: consequently the plant is the PLANTS) gametophyte, and the sporophyte is represented only by the oospore, so that the life-history resembles that of Nemalion. It is probable that this conclusion is generally true of the whole group; at any rate of those forms (Desmids, Spirogyra, Oedogonium, Chara) which have been more or less investigated. Turning to the Fungi, somewhat similar results have been obtained in the few forms that have been studied from this point of view. In the sexual Ascomycetes it appears (Harper) that meiosis takes place in the ascocarp just before the development of the spores, so that the life-history essentially resembles that of Nemalion. Again, in certain Uredineae, having an aecidium-stage and a teleutospore-stage, which is apparently a sexual process has been observed (Blackman, Christman) which is described in the section on Abnormalities of Reproduction, and the life-history is as follows. The sexual act having taken place, a row of aecidiospores is developed in the aecidium, each of which contains two conjugate nuclei derived from the sexual nuclei. The mycelium developed from the aecidiospore, as well as the uredospores and the teleutospores that it bears, shows two conjugate nuclei. When, however, the teleutospore is about to germinate, the two nuclei fuse (thus completing the sexual act) and meiosis takes place. As a result the promycelium developed from the teleutospore, and the sporidia that it produces, are uninucleate: so are also the mycelium developed from the sporidium, and the female organs (archicarps) borne upon it. Hence the limits of the sporophyte are the aecidiospore and the teleutospore: those of the gametophyte, the teleutospore and the aecidiospore. Similar observations have been made upon other Uredineae with a more contracted life-history. Phragmidium Potentillaecanadensis is a rust that has no aecidium-stage: consequently the primary uredospores are borne by the mycelium produced on infection of the host by a sporidium. It has been observed (Christman) that the sporogenous hyphae fuse in pairs, suggesting a sexual act; then the primary uredospores are developed in rows from the fused pairs of hyphae which thus behave as sexual organs (archicarps), and each such uredospore contains two conjugate nuclei. Although the research has not been carried beyond this point, it may be inferred that in this case, as in the preceding, nuclear fusion and meiosis take place in the teleutospore. Here the sporophyte is represented by the uredo-form. Finally, in some of the fungi in which no sexual organs have yet been discovered, this method of investigation has made it probable that some kind of sexual act takes place nevertheless. Thus in the Uredine Puccinia malvacearum, which has only teleutospore- and sporidium-stages, it has been observed (Black-man) that the formation of the teleutospores is preceded by a binucleate condition of the hyphae. The same idea is suggested by the binucleate basidia of the Basidiomycetes, which correspond to the teleutospores of the Uredineae. The life-histories sketched in the preceding paragraphs show that one of the complexities met with in the Thallophyta is that meiosis does not always take place at the same point in the life-history. In the higher plants the incidence of meiosis is generally, though not absolutely, constant: it may be stated as a rule that in the Bryophyta, Pteridophyta and Phanerogams it takes place in the spore-mother-cells. In the Thallophyta this rule does not hold. In some of them, it is true, meiosis immediately precedes, as in the higher plants, the formation of certain spores, the tetraspores (Dictyotaceae, Polysiphonia), the teleutospores (Uredineae): but in others it immediately precedes the development of the sexual organs (Fucaceae), or follows more or less directly upon the sexual act (Green Algae, Nemalion, Ascomycetes). The life-history of most Thallophyta is further complicated by the capacity of the gametophyte of the sporophyte to repro-duce themselves by cells termed gonidia, a capacity that is wholly lacking in the higher plants. The karyology of gonidia has not yet been sufficiently investigated: but when, as in the Green Algae and the Oomycetous Fungi, the gonidia are developed125 by and reproduce the gametophyte, it may be inferred that they, like the gametophyte, are haploid. One case, at any rate, of the reproduction of the sporophyte by gonidia is fully known, that of the Uredineae just described, in which the uredoform, which is a phase of the sporophyte, is reproduced by the uredo-spores which are binucleate, that is diploid, and may be distinguished as diplogonidia. In any case the result is that whereas in the higher plants each of the alternating generations occurs but once in the life-history, in these Thallophyta the life-history may include a succession of gametophytic or of sporophytic forms This is, in fact, a distinguishing feature of the group. The higher plants present a regular alternation of generations: whereas, in the Thallophyta, though they probably all present some kind of alternation of generations, yet it is irregular in the various ways and for the various reasons mentioned above Sufficient information has been given in the preceding pages to render possible the consideration of the origin of alternation of generations. To begin quite at the beginning, it may be assumed that the primitive form of reproduction was purely vegetative, merely division of the unicellular organism when it had attained the limits of its own growth. Following on this came reproduction by a gonidium: that is, the protoplasm of the cell, at the end of its vegetative life, became quiescent, surrounded itself with a proper wall, or was set free as a motile ciliated cell, having in some unexplained way become capable of originating a new course of life (rejuvenescence) on germination. Then, as can be well traced in the Brown and Green Algae (see ALGAE), these primitive reproductive cells (gonidia) began to fuse in pairs: in other words, they gradually became sexual. This stage can still be observed in some of these Algae (e.g. Ulothrix, Ectocarpus) where the zoospores (gonidia) may either germinate independently, or fuse in pairs to form a zygote. Gradually the sexuality of these cells became more pronounced: losing the capacity for independent germination, they acquired the external characters of more or less differentiated sexual cells, and the gametangia producing them developed into male and female sexual organs. But this advancing sexual differentiation did not necessarily deprive the plant of the primitive mode of propagation: the sexual organism still retained the faculty of reproduction by gonidia. The loss of this faculty only came with higher development: it is entirely wanting in some of the higher Thallophyta (e.g. Fucaceae, Characeae), and in all plants above them in the evolutionary series. With the introduction of the sexual act, a new kind of reproductive cell made its appearance, the zygote. This cell, as already explained, differs from other kinds of spores and from the sexual cells, in that its necleus is diploid; and with it the sporophyte (diplophyte) was introduced into the life-history. It has been mentioned that in some plants (e.g. Green Algae) the zygote is all that there is to represent the sporophyte, giving rise, or germination and after meiosis, to one or more spores. Passing to the Bryophyta, in the simpler forms (e.g. Riccia), the zygote develops into a multicellular capsule (sporogonium); and in the higher forms into a more elaborate sporogonium, producing many spores. In the Pteridophyta and the Phanerogams, the zygote gives rise to the highly developed sporophytic plant. Thus the evolution of the sporophyte can be traced from the unicellular zygote, gradually increasing in bulk and in in-dependence until it becomes the equal of the gametophyte (e.g. in Dictyota and Polysiphonia), and eventually far surpasses it (Pteridophyta, Phanerogams). Moreover, the increase in size was attended by the gradual limitation of spore-production to certain parts only, the rest of the tissues being vegetative, assuming the form of stems, leaves, &c. These facts have been formulated in the theory of " progressive sterilization " (Bower), which states that the sporophytic form of the higher plants has been evolved from the simple, entirely fertile, sporophyte of the lower, by the gradually increasing development of the sterile vegetative tissue at the expense of the sporogenous, accompanied by increase in total bulk and in morphological and histological differentiation. In connexion with the study of the evolution of the sporophyte, the question arose as to its morphological significance; whether it is to be regarded as a modified form of the gametophyte, or as an altogether new form intercalated in the life-history: in other words, whether the alternation is" homologous "or" antithetic." In certain plants there is a succession of forms which are undoubtedly homologous: for instance, in Coleochaete where a succession of individuals without sexual organs is produced by zoospores (gonidia). The main fact that has been established is that the sporophyte, from the simple zygote of the Thallophyta to the spore-bearing plant of the Phanerogams, is characterized by its diploid nuclei; that it is a diplophyte, in contrast to the haplophytic gametophyte. Were these nuclear characters absolutely universal, there could be no question but that the sporophyte is an altogether new antithetic form, and not an homologous generation. But certain exceptions to the rule have been detected, which are described under Abnormalities of Reproduction: at present it will suffice to say that such things as a diploid gametophyte and a haploid sporophyte have been observed in certain ferns. It can only be inferred that alternation of generations is not absolutely dependent upon the periodic halving in meiosis and the subsequent doubling by a sexual act, of the number of chromosomes in the nuclei, though the two sets of phenomena usually coincide. It must not, however, be overlooked that these exceptional cases occur in plants presenting an abnormal life-history: the fact remains that where there is both normal spore-formation with meiosis, and a subsequent sexual act, the haploid form is the gametophyte, the diploid the sporophyte. But the actual observation of a haploid sporophyte and of a diploid gametophyte makes it clear that however generally useful the nuclear characters may be in the distinction of sporophyte and gametophyte, they do not afford an absolute criterion, and therefore their value in determining homologies is debatable. IV. Abnormalities of Reproduction. In what may be regarded as the type of normal life-history, the transition from the one generation to the other is marked by definite processes: there is the meiotic development of spores by the sporophyte, and the sexual production of a zygote, or something analogous to it, by the gametophyte. But it has been mentioned in the preceding pages that the transition may, in certain cases, be effected in other ways, which may be regarded as abnormal, though they are constant enough in the plants in which they occur, in fact as manifestations of reproductive degeneration. In the first place, the sporophyte may be developed either after an abnormal sexual act, or without any preceding sexual act at all, a condition known as apogamy. In the second, the gametophyte may be developed otherwise than from a post- meiotic spore, a condition known as apospory.

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