Alternation of generations (also known as metagenesis or heterogenesis) is the predominant type of
life cycle
Life cycle, life-cycle, or lifecycle may refer to:
Science and academia
*Biological life cycle, the sequence of life stages that an organism undergoes from birth to reproduction ending with the production of the offspring
* Life-cycle hypothesis ...
in
plant
Plants are predominantly photosynthetic eukaryotes of the kingdom Plantae. Historically, the plant kingdom encompassed all living things that were not animals, and included algae and fungi; however, all current definitions of Plantae exclud ...
s and
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
. It consists of a
multicellular
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism.
All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- ...
haploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
sexual phase, the
gametophyte
A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...
, which has a single set of
chromosome
A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s alternating with a multicellular
diploid
Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
asexual phase, the
sporophyte
A sporophyte () is the diploid multicellular stage in the life cycle of a plant or alga which produces asexual spores. This stage alternates with a multicellular haploid gametophyte phase.
Life cycle
The sporophyte develops from the zygote pr ...
which has two sets of chromosomes.
A mature sporophyte produces haploid
spore
In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
s by
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
, a process which reduces the number of chromosomes to half, from two sets to one. The resulting haploid spores germinate and grow into multicellular haploid gametophytes. At maturity, a gametophyte produces
gamete
A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s by
mitosis
In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
, the normal process of cell division in eukaryotes, which maintains the original number of chromosomes. Two haploid gametes (originating from different organisms of the same
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
or from the same organism)
fuse
Fuse or FUSE may refer to:
Devices
* Fuse (electrical), a device used in electrical systems to protect against excessive current
** Fuse (automotive), a class of fuses for vehicles
* Fuse (hydraulic), a device used in hydraulic systems to protect ...
to produce a diploid
zygote
A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism.
In multicellula ...
, which divides repeatedly by mitosis, developing into a multicellular diploid sporophyte. This cycle, from gametophyte to sporophyte (or equally from sporophyte to gametophyte), is the way in which all land plants and most
algae
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mic ...
undergo
sexual reproduction
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
.
The relationship between the sporophyte and gametophyte phases varies among different groups of plants. In the majority of
alga
Algae (; singular alga ) is an informal term for a large and diverse group of photosynthetic eukaryotic organisms. It is a polyphyletic grouping that includes species from multiple distinct clades. Included organisms range from unicellular mi ...
e, the sporophyte and gametophyte are separate independent organisms, which may or may not have a similar appearance. In
liverwort
The Marchantiophyta () are a division of non-vascular land plants commonly referred to as hepatics or liverworts. Like mosses and hornworts, they have a gametophyte-dominant life cycle, in which cells of the plant carry only a single set of g ...
s,
moss
Mosses are small, non-vascular flowerless plants in the taxonomic division Bryophyta (, ) '' sensu stricto''. Bryophyta (''sensu lato'', Schimp. 1879) may also refer to the parent group bryophytes, which comprise liverworts, mosses, and hor ...
es and
hornwort
Hornworts are a group of non-vascular Embryophytes (land plants) constituting the division Anthocerotophyta (). The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, hornworts have a ...
s, 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
Vascular plants (), also called tracheophytes () or collectively Tracheophyta (), form a large group of land plants ( accepted known species) that have lignified tissues (the xylem) for conducting water and minerals throughout the plant. They al ...
the gametophyte is less well developed than the sporophyte, although their
Devonian
The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...
ancestors had gametophytes and sporophytes of approximately equivalent complexity. In
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s the gametophyte is a small flattened
autotrophic
An autotroph or primary producer is an organism that produces complex organic compounds (such as carbohydrates, fats, and proteins) using carbon from simple substances such as carbon dioxide,Morris, J. et al. (2019). "Biology: How Life Works", ...
prothallus
A prothallus, or prothallium, (from Latin ''pro'' = forwards and Greek ''θαλλος'' (''thallos'') = twig) is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young ...
on which the young sporophyte is briefly dependent for its nutrition. In
flowering plant
Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s, 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. (Some insects have a sex-determining system whereby haploid males are produced from unfertilized eggs; however females produced from fertilized eggs are diploid.)
Life cycles of plants and algae with alternating haploid and diploid multicellular stages are referred to as diplohaplontic (the equivalent terms haplodiplontic, diplobiontic and dibiontic are also in use, as is describing such an organism as having a diphasic
ontogeny
Ontogeny (also ontogenesis) is the origination and development of an organism (both physical and psychological, e.g., moral development), usually from the time of fertilization of the egg to adult. The term can also be used to refer to the stu ...
). 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 these forms are free-living. In some species, such as the alga ''
Ulva lactuca
''Ulva lactuca'', also known by the common name sea lettuce, is an edible green alga in the family Ulvaceae. It is the type species of the genus ''Ulva''. A synonym is ''U. fenestrata'', referring to its "windowed" or "holed" appearance.
Descrip ...
'', 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
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
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 Bryophyta s.l. are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. Bryophyta s.s. consists of the mosses only. They are characteristically limited i ...
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 plant
A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
s, 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 (just three cells in many cases). Here the notion of two generations is less obvious; as Bateman & Dimichele say " orophyte 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 (sexual vs. asexual, gametophyte vs. sporophyte, haploid vs. diploid, etc.).
Initially,
Chamisso
Adelbert von Chamisso (; 30 January 178121 August 1838) was a German poet and botanist, author of ''Peter Schlemihl'', a famous story about a man who sold his shadow. He was commonly known in French as Adelbert de Chamisso (or Chamissot) de Bonc ...
and Steenstrup described the succession of differently organized generations (sexual and asexual) in animals as "alternation of generations", while studying the development of
tunicate
A tunicate is a marine invertebrate animal, a member of the subphylum Tunicata (). It is part of the Chordata, a phylum which includes all animals with dorsal nerve cords and notochords (including vertebrates). The subphylum was at one time ca ...
trematode
Trematoda is a class of flatworms known as flukes. They are obligate internal parasites with a complex life cycle requiring at least two hosts. The intermediate host, in which asexual reproduction occurs, is usually a snail. The definitive host ...
animals. This phenomenon is also known as
heterogamy
Heterogamy is a term applied to a variety of distinct phenomena in different scientific domains. Usually having to do with some kind of difference, "hetero", in reproduction, "gamy". See below for more specific senses.
Science Reproductive biolog ...
. 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
Wilhelm Friedrich Benedikt Hofmeister (18 May 1824 – 12 January 1877) was a German biologist and botanist. He "stands as one of the true giants in the history of biology and belongs in the same pantheon as Darwin and Mendel." He was lar ...
demonstrated the morphological alternation of generations in plants, between a spore-bearing generation (sporophyte) and a gamete-bearing generation (gametophyte). By that time, a debate emerged focusing on the origin of the asexual generation of land plants (i.e., the sporophyte) and is conventionally characterized as a conflict between theories of antithetic ( Čelakovský, 1874) and homologous (
Pringsheim Pringsheim is a Jewish Silesian surname. Notable people with the surname include:
* Alfred Pringsheim (1850–1941), mathematician, father-in-law of writer Thomas Mann
* Ernst Pringsheim Sr. (1859–1917), German physicist
* Ernst Pringsheim Jr. ...
, 1876) alternation of generations. Čelakovský coined the words sporophyte and gametophyte.
Eduard Strasburger
Eduard Adolf Strasburger (1 February 1844 – 18 May 1912) was a Polish-German professor and one of the most famous botanists of the 19th century. He discovered mitosis in plants.
Life
Eduard Strasburger was born in Warsaw, Congress Poland, the ...
(1874) 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
Red algae, or Rhodophyta (, ; ), are one of the oldest groups of eukaryotic algae. The Rhodophyta also comprises one of the largest phyla of algae, containing over 7,000 currently recognized species with taxonomic revisions ongoing. The majority ...
, the same nuclear phase may correspond to two diverse morphological generations. In some
fern
A fern (Polypodiopsida or Polypodiophyta ) is a member of a group of vascular plants (plants with xylem and phloem) that reproduce via spores and have neither seeds nor flowers. The polypodiophytes include all living pteridophytes except t ...
s 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. 2''n'' chromosomes in total.
* The single-celled diploid zygote germinates, dividing by the normal process (
mitosis
In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is mainta ...
), which maintains the number of chromosomes at 2''n''. The result is a multi-cellular diploid organism, called the ''sporophyte'' (because at maturity it produces spores).
* When it reaches maturity, the sporophyte produces one or more sporangia (singular: sporangium) which are the organs that produce diploid spore mother cells (sporocytes). These divide by a special process (
meiosis
Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
) 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 (mitosis), which maintains the number of chromosomes at ''n''. The result is a multi-cellular haploid organism, called the ''gametophyte'' (because it produces gametes at maturity).
* When it reaches maturity, the gametophyte produces one or more gametangia (singular: gametangium) 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.
The 'alternation of generations' in the life cycle is thus between a diploid (2''n'') generation of sporophytes and a haploid (''n'') generation of gametophytes.
The situation is quite different from that in animals, where the fundamental process is that a diploid (2''n'') individual produces haploid (''n'') gametes by meiosis. In animals, spores (i.e. haploid cells which are able to undergo mitosis) are not produced, so there is no asexual multi-cellular generation. Some insects have haploid males that develop from unfertilized eggs, but the females are all diploid.
Variations
The diagram shown above is a good representation of the life cycle of some multi-cellular algae (e.g. the genus ''
Cladophora
''Cladophora'' is a genus of reticulated filamentous Ulvophyceae (green algae). The genus ''Cladophora'' contains many species that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is a ...
'') 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'' (homomorphy or isomorphy). Filamentous algae of the genus ''
Cladophora
''Cladophora'' is a genus of reticulated filamentous Ulvophyceae (green algae). The genus ''Cladophora'' contains many species that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is a ...
'', 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'' (heteromorphy or anisomorphy).
*** ''Dominant gametophyte'' (gametophytic). 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.
*** ''Dominant sporophyte'' (sporophytic). 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
A gametophyte () is one of the two alternation of generations, alternating multicellular organism, multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has on ...
is even more reduced (at the minimum to only three cells), 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: " orophyte 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'' (isogamy). Like other species of ''
Cladophora
''Cladophora'' is a genus of reticulated filamentous Ulvophyceae (green algae). The genus ''Cladophora'' contains many species that are very hard to tell apart and classify, mainly because of the great variation in their appearances, which is a ...
'', ''C. callicoma'' has flagellated gametes which are identical in appearance and ability to move.
** ''Gametes of two distinct sizes'' (anisogamy).
*** ''Both of similar motility''. Species of ''
Ulva
Ulva (; gd, Ulbha) is a small island in the Inner Hebrides of Scotland, off the west coast of Mull. It is separated from Mull by a narrow strait, and connected to the neighbouring island of Gometra by a bridge. Much of the island is formed from ...
'', 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'' (oogamy). The larger sessile megagametes are eggs (ova), and smaller motile