Alternation of generations
Alternation of generations is the type of life cycle that occurs in those plants and algae in the Archaeplastida and the Heterokontophyta that have distinct haploid sexual and diploid asexual stages. In these groups, a multicellular haploid gametophyte with n chromosomes alternates with a multicellular diploid sporophyte with 2n chromosomes, made up of n pairs. A mature sporophyte produces haploid spores by meiosis, a process which reduces the number of chromosomes to half, from 2n to n.
The haploid spores germinate and grow into a haploid gametophyte. At maturity, the gametophyte produces gametes by mitosis, which does not alter the number of chromosomes. Two gametes fuse to produce a diploid zygote, which develops into a diploid sporophyte. This cycle, from gametophyte to sporophyte, is the way in which all land plants and many algae undergo sexual reproduction.
The relationship between the sporophyte and gametophyte varies among different groups of plants. In those algae which have alternation of generations, the sporophyte and gametophyte are separate independent organisms, which may or may not have a similar appearance. In liverworts, mosses and hornworts, the sporophyte is less well developed than the gametophyte and is largely dependent on it. Although moss and hornwort sporophytes can photosynthesise, they require additional photosynthate from the gametophyte to sustain growth and spore development and depend on it for supply of water, mineral nutrients and nitrogen. By contrast, in all modern vascular plants the gametophyte is less well developed than the sporophyte, although their Devonian ancestors had gametophytes and sporophytes of approximately equivalent complexity. In ferns the gametophyte is a small flattened autotrophic prothallus on which the young sporophyte is briefly dependent for its nutrition. In flowering plants, the reduction of the gametophyte is much more extreme; it consists of just a few cells which grow entirely inside the sporophyte.
Animals develop differently. They produce haploid gametes. No haploid spores capable of dividing are produced, so generally there is no multicellular haploid phase.
Life cycles of plants and algae with alternating haploid and diploid multicellular stages are referred to as diplohaplontic. Life cycles, such as those of animals, in which there is only a diploid multicellular stage are referred to as diplontic. Life cycles in which there is only a haploid multicellular stage are referred to as haplontic.
Definition
Alternation of generations is defined as the alternation of multicellular diploid and haploid forms in the organism's life cycle, regardless of whether or not these forms are free-living. In some species, such as the alga Ulva lactuca, the diploid and haploid forms are indeed both free-living independent organisms, essentially identical in appearance and therefore said to be isomorphic. The free-swimming, haploid gametes form a diploid zygote which germinates into a multicellular diploid sporophyte. The sporophyte produces free-swimming haploid spores by meiosis that germinate into haploid gametophytes.However, in some other groups, either the sporophyte or the gametophyte is very much reduced and is incapable of free living. For example, in all bryophytes the gametophyte generation is dominant and the sporophyte is dependent on it. By contrast, in all modern vascular land plants the gametophytes are strongly reduced, although the fossil evidence indicates that they were derived from isomorphic ancestors. In seed plants, the female gametophyte develops totally within the sporophyte, which protects and nurtures it and the embryonic sporophyte that it produces. The pollen grains, which are the male gametophytes, are reduced to only a few cells. Here the notion of two generations is less obvious; as Bateman & Dimichele say "porophyte and gametophyte effectively function as a single organism". The alternative term 'alternation of phases' may then be more appropriate.
History
Debates about alternation of generations in the early twentieth century can be confusing because various ways of classifying "generations" co-exist.Initially, Chamisso and Steenstrup described the succession of differently organized generations in animals as "alternation of generations", while studying the development of tunicates, cnidarians and trematode animals. This phenomenon is also known as heterogamy. Presently, the term "alternation of generations" is almost exclusively associated with the life cycles of plants, specifically with the alternation of haploid gametophytes and diploid sporophytes.
Wilhelm Hofmeister demonstrated the morphological alternation of generations in plants, between a spore-bearing generation and a gamete-bearing generation. By that time, a debate emerged focusing on the origin of the asexual generation of land plants and is conventionally characterized as a conflict between theories of antithetic and homologous alternation of generations. Čelakovský coined the words sporophyte and gametophyte.
Eduard Strasburger discovered the alternation between diploid and haploid nuclear phases, also called cytological alternation of nuclear phases. Although most often coinciding, morphological alternation and nuclear phases alternation are sometimes independent of one another, e.g., in many red algae, the same nuclear phase may correspond to two diverse morphological generations. In some ferns which lost sexual reproduction, there is no change in nuclear phase, but the alternation of generations is maintained.
Alternation of generations in plants
Fundamental elements
The diagram above shows the fundamental elements of the alternation of generations in plants. The many variations found in different groups of plants are described by use of these concepts later in the article. Starting from the right of the diagram, the processes involved are as follows:- Two single-celled haploid gametes, each containing n unpaired chromosomes, fuse to form a single-celled diploid zygote, which now contains n pairs of chromosomes, i.e. 2n chromosomes in total.
- The single-celled diploid zygote germinates, dividing by the normal process, which maintains the number of chromosomes at 2n. The result is a multi-cellular diploid organism, called the sporophyte.
- When it reaches maturity, the sporophyte produces one or more sporangia which are the organs that produce diploid spore mother cells. These divide by a special process that reduces the number of chromosomes by a half. This initially results in four single-celled haploid spores, each containing n unpaired chromosomes.
- The single-celled haploid spore germinates, dividing by the normal process, which maintains the number of chromosomes at n. The result is a multi-cellular haploid organism, called the gametophyte.
- When it reaches maturity, the gametophyte produces one or more gametangia which are the organs that produce haploid gametes. At least one kind of gamete possesses some mechanism for reaching another gamete in order to fuse with it.
'' with a descendant sporophyte beginning to grow from it.
The situation is quite different from that in animals, where the fundamental process is that a diploid individual produces haploid gametes by meiosis. Spores are not produced, so neither is an asexual multi-cellular generation that alternates with a sexual multi-cellular generation.
Variations
The diagram shown above is a good representation of the life cycle of some multi-cellular algae which have sporophytes and gametophytes of almost identical appearance and which do not have different kinds of spores or gametes.However, there are many possible variations on the fundamental elements of a life cycle which has alternation of generations. Each variation may occur separately or in combination, resulting in a bewildering variety of life cycles. The terms used by botanists in describing these life cycles can be equally bewildering. As Bateman and Dimichele say " the alternation of generations has become a terminological morass; often, one term represents several concepts or one concept is represented by several terms."
Possible variations are:
- Relative importance of the sporophyte and the gametophyte.
- * Equal.
Filamentous algae of the genus Cladophora, which are predominantly found in fresh water, have diploid sporophytes and haploid gametophytes which are externally indistinguishable. No living land plant has equally dominant sporophytes and gametophytes, although some theories of the evolution of alternation of generations suggest that ancestral land plants did. - * Unequal., a moss.
- ** Dominant gametophyte.
In liverworts, mosses and hornworts, the dominant form is the haploid gametophyte. The diploid sporophyte is not capable of an independent existence, gaining most of its nutrition from the parent gametophyte, and having no chlorophyll when mature., a fern. - ** Dominant sporophyte.
In ferns, both the sporophyte and the gametophyte are capable of living independently, but the dominant form is the diploid sporophyte. The haploid gametophyte is much smaller and simpler in structure. In seed plants, the gametophyte is even more reduced, gaining all its nutrition from the sporophyte. The extreme reduction in the size of the gametophyte and its retention within the sporophyte means that when applied to seed plants the term 'alternation of generations' is somewhat misleading: "porophyte and gametophyte effectively function as a single organism". Some authors have preferred the term 'alternation of phases'. - Differentiation of the gametes.
- * Both gametes the same.
Like other species of Cladophora, C. callicoma has flagellated gametes which are identical in appearance and ability to move. - * Gametes of two distinct sizes.
- ** Both of similar motility.
Species of Ulva, the sea lettuce, have gametes which all have two flagella and so are motile. However they are of two sizes: larger 'female' gametes and smaller 'male' gametes. - ** One large and sessile, one small and motile. The larger sessile megagametes are eggs, and smaller motile microgametes are sperm. The degree of motility of the sperm may be very limited but all are able to move towards the sessile eggs. When the sperm and eggs are produced in different kinds of gametangia, the sperm-producing ones are called antheridia and the egg-producing ones archegonia. with sporophytes growing from the remains of archegonia.
- *** Antheridia and archegonia occur on the same gametophyte, which is then called monoicous.
The liverwort Pellia epiphylla has the gametophyte as the dominant generation. It is monoicous: the small reddish sperm-producing antheridia are scattered along the midrib while the egg-producing archegonia grow nearer the tips of divisions of the plant. - *** Antheridia and archegonia occur on different gametophytes, which are then called dioicous.
The moss Mnium hornum has the gametophyte as the dominant generation. It is dioicous: male plants produce only antheridia in terminal rosettes, female plants produce only archegonia in the form of stalked capsules. Seed plant gametophytes are also dioicous. However, the parent sporophyte may be monoecious, producing both male and female gametophytes or dioecious, producing gametophytes of one gender only. Seed plant gametophytes are extremely reduced in size; the archegonium consists only of a small number of cells, and the entire male gametophyte may be represented by only two cells. - Differentiation of the spores.
- * All spores the same size.
Horsetails have spores which are all of the same size. - * Spores of two distinct sizes : larger megaspores and smaller microspores. When the two kinds of spore are produced in different kinds of sporangia, these are called megasporangia and microsporangia. A megaspore often develops at the expense of the other three cells resulting from meiosis, which abort.
- ** Megasporangia and microsporangia occur on the same sporophyte, which is then called monoecious.
Most flowering plants fall into this category. Thus the flower of a lily contains six stamens which produce microspores which develop into pollen grains, and three fused carpels which produce integumented megasporangia each of which produces a megaspore which develops inside the megasporangium to produce the megagametophyte. In other plants, such as hazel, some flowers have only stamens, others only carpels, but the same plant has both kinds of flower and so is monoecious. - ** Megasporangia and microsporangia occur on different sporophytes, which are then called dioecious.
An individual tree of the European holly produces either 'male' flowers which have only functional stamens producing microspores which develop into pollen grains or 'female' flowers which have only functional carpels producing integumented megasporangia that contain a megaspore that develops into a multicellular megagametophyte.
A complex life cycle
The diagram shows the alternation of generations in a species which is heteromorphic, sporophytic, oogametic, dioicous, heterosporic and dioecious. A seed plant example might be a willow tree. Starting in the centre of the diagram, the processes involved are:- An immobile egg, contained in the archegonium, fuses with a mobile sperm, released from an antheridium. The resulting zygote is either 'male' or 'female'.
- * A 'male' zygote develops by mitosis into a microsporophyte, which at maturity produces one or more microsporangia. Microspores develop within the microsporangium by meiosis.
In a willow the zygote first develops into an embryo microsporophyte within the ovule. At maturity, these structures become the seed. Later the seed is shed, germinates and grows into a mature tree. A 'male' willow tree produces flowers with only stamens, the anthers of which are the microsporangia. - * Microspores germinate producing microgametophytes; at maturity one or more antheridia are produced. Sperm develop within the antheridia.
In a willow, microspores are not liberated from the anther, but develop into pollen grains within it. The whole pollen grain is moved to an ovule, where a sperm is produced which moves down a pollen tube to reach the egg. - * A 'female' zygote develops by mitosis into a megasporophyte, which at maturity produces one or more megasporangia. Megaspores develop within the megasporangium; typically one of the four spores produced by meiosis gains bulk at the expense of the remaining three, which disappear.
'Female' willow trees produce flowers with only carpels. - * Megaspores germinate producing megagametophytes; at maturity one or more archegonia are produced. Eggs develop within the archegonia.
The carpels of a willow produce ovules, megasporangia enclosed in integuments. Within each ovule, a megaspore develops by mitosis into a megagametophyte. An archegonium develops within the megagametophyte and produces an egg. The whole of the gametophytic 'generation' remains within the protection of the sporophyte except for pollen grains.Life cycles of different plant groups
Alternation of generations occurs in almost all multicellular red and green algae, both freshwater forms and seaweeds. In most, the generations are homomorphic and free-living. Some species of red algae have a complex triphasic alternation of generations, in which there is a gametophyte phase and two distinct sporophyte phases. For further information, see Red algae: Reproduction.
Land plants all have heteromorphic alternation of generations, in which the sporophyte and gametophyte are distinctly different. All bryophytes, i.e. liverworts, mosses and hornworts, have the gametophyte generation as the most conspicuous. As an illustration, consider a monoicous moss. Antheridia and archegonia develop on the mature plant. In the presence of water, the biflagellate sperm from the antheridia swim to the archegonia and fertilisation occurs, leading to the production of a diploid sporophyte. The sporophyte grows up from the archegonium. Its body comprises a long stalk topped by a capsule within which spore-producing cells undergo meiosis to form haploid spores. Most mosses rely on the wind to disperse these spores, although Splachnum sphaericum is entomophilous, recruiting insects to disperse its spores. For further information, see Liverwort: Life cycle, Moss: Life cycle, Hornwort: Life cycle.
In ferns and their allies, including clubmosses and horsetails, the conspicuous plant observed in the field is the diploid sporophyte. The haploid spores develop in sori on the underside of the fronds and are dispersed by the wind. If conditions are right, a spore will germinate and grow into a rather inconspicuous plant body called a prothallus. The haploid prothallus does not resemble the sporophyte, and as such ferns and their allies have a heteromorphic alternation of generations. The prothallus is short-lived, but carries out sexual reproduction, producing the diploid zygote that then grows out of the prothallus as the sporophyte. For further information, see Fern: Life cycle.
In the spermatophytes, the seed plants, the sporophyte is the dominant multicellular phase; the gametophytes are strongly reduced in size and very different in morphology. The entire gametophyte generation, with the sole exception of pollen grains, is contained within the sporophyte. The life cycle of a dioecious flowering plant, the willow, has been outlined in some detail in an earlier section. The life cycle of a gymnosperm is similar. However, flowering plants have in addition a phenomenon called 'double fertilization'. Two sperm nuclei from a pollen grain, rather than a single sperm, enter the archegonium of the megagametophyte; one fuses with the egg nucleus to form the zygote, the other fuses with two other nuclei of the gametophyte to form 'endosperm', which nourishes the developing embryo. For further information, see Double fertilization.