Antheridiogens are a class of chemicals secreted by fern gametophytes [1] that have "been shown to influence production of male gametangia and thus mating systems in a large number of terrestrial fern species". [2] Antheridiogens are only observed in homosporous fern species, as all gametophytes are potentially bisexual (have the ability to produce both archegonia and antheridia). [3]
The first study regarding antheridiogen was published by Walter Döpp in 1950. [4] In this article, he explains the discovery of a molecule, which he titled "A-substanz", that caused premature formation of antheridia when agar media was reused after cultivation of Pteridium aquilinum. [5] A majority of the studies regarding antheridiogen were done by two researchers, Ulrich Näf and H. Schraudolf. [6]
The way in which antheridiogen determines sex in ferns is a "spatiotemporally split gibberellin synthesis pathway". [7] Gibberellin is a group of hormones that control plant processes. [8] In the first step of this process, gametophytes, or prothalli, express gibberellin (GA) specific genes, which produces a GA intermediate molecule that is then secreted into the external environment. In the second step, antheridiogens are taken up by neighboring gametophytes in the colony and undergoes a series of molecular changes that allow it to finally induce or suppress formation of antheridia or archegonia. This helps regulate the sex ratio of the colony. [7]
The timing with which antheridiogen affects the gender of growing gametophytes is still under study. One theory states that "the spores that germinate first develop as hermaphrodites and secrete antheridiogen, while those that germinate later or develop more slowly become male under the influence of the secreted antheridiogen". [9] Depending on the ratio of males to hermaphrodites, either outcrossing or inbreeding is selected for by the population.
Studies performed on Ceratopteris richardii have proven that a growing gametophyte is only able to respond to its own antheridiogen for a brief period of time in very early stages of growth, and that if exposure is removed, undifferentiated cells can revert from male to hermaphrodite. [9] The reversibility of male expression proves that levels of antheridiogen must be maintained in order to keep male expression from disappearing.
Antheridiogen has also been shown to allow spores that are grown in complete darkness to grow. [5] Spores that are buried underground, if reached by antheridiogen, can form gametophytes that reach the surface. Or, they form a small amount of antheridia, and the sperm produced can reach the female gametophytes above ground.
The way in which the sex of each individual is determined is a form of environmental sex determination (ESD). In contrast to chromosomal sex determination, sex determination through ESD is extremely flexible, allowing for sex changes throughout the individual's life in response to colony or environmental changes. One study performed on the fern species Woodwardia radicans found that sex expression of mature gametophytes is dependent on stress conditions. [10] In good conditions, gametophytes reached sexual maturity at a larger size and became females and then bisexuals, whereas in stressful conditions, gametophytes reached sexual maturity at a smaller size and became males.
The idea of an antheridiogen is very similar to that of pheromones, which exist in some mammals. Although mammalian sex determination is decided by chromosomes, chemicals called pheromones are released, detected by the olfactory system, and can control social behaviors and hormone levels. [11] The presence of human pheromones remains a long-contested topic, as there have not been any studies that completely prove the existence of pheromones or of a possible pathway of detection.
A gametophyte is one of the two alternating multicellular phases in the life cycles of plants and algae. It is a haploid multicellular organism that develops from a haploid spore that has one set of chromosomes. The gametophyte is the sexual phase in the life cycle of plants and algae. It develops sex organs that produce gametes, haploid sex cells that participate in fertilization to form a diploid zygote which has a double set of chromosomes. Cell division of the zygote results in a new diploid multicellular organism, the second stage in the life cycle known as the sporophyte. The sporophyte can produce haploid spores by meiosis that on germination produce a new generation of gametophytes.
A fern is a member of a group of vascular plants that reproduce via spores and have neither seeds nor flowers. They differ from mosses and other bryophytes by being vascular, i.e., having specialized tissues that conduct water and nutrients and in having life cycles in which the branched sporophyte is the dominant phase. Ferns have complex leaves called megaphylls, that are more complex than the microphylls of clubmosses. Most ferns are leptosporangiate ferns. They produce coiled fiddleheads that uncoil and expand into fronds. The group includes about 10,560 known extant species. Ferns are defined here in the broad sense, being all of the Polypodiopsida, comprising both the leptosporangiate (Polypodiidae) and eusporangiate ferns, the latter group including horsetails or scouring rushes, whisk ferns, marattioid ferns, and ophioglossoid ferns.
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.
Bracken (Pteridium) is a genus of large, coarse ferns in the family Dennstaedtiaceae. Ferns (Pteridophyta) are vascular plants that have alternating generations, large plants that produce spores and small plants that produce sex cells. Brackens are noted for their large, highly divided leaves. They are found on all continents except Antarctica and in all environments except deserts, though their typical habitat is moorland. The genus probably has the widest distribution of any fern in the world.
Bryophytes are a proposed taxonomic division containing three groups of non-vascular land plants (embryophytes): the liverworts, hornworts and mosses. They are characteristically limited in size and prefer moist habitats although they can survive in drier environments. The bryophytes consist of about 20,000 plant species. Bryophytes produce enclosed reproductive structures, but they do not produce flowers or seeds. They reproduce by spore formation. Though bryophytes were considered a paraphyletic group in recent years, almost all of the most recent phylogenetic evidence supports the monophyly of this group, as originally classified by Wilhelm Schimper in 1879. The term "bryophyte" comes from Greek βρύον, bryon "tree-moss, oyster-green" and φυτόν, phyton "plant".
A sporophyte is the diploid multicellular stage in the life cycle of a plant or alga. It develops from the zygote produced when a haploid egg cell is fertilized by a haploid sperm and each sporophyte cell therefore has a double set of chromosomes, one set from each parent. All land plants, and most multicellular algae, have life cycles in which a multicellular diploid sporophyte phase alternates with a multicellular haploid gametophyte phase. In the seed plants, the largest groups of which are the gymnosperms and flowering plants (angiosperms), the sporophyte phase is more prominent than the gametophyte, and is the familiar green plant with its roots, stem, leaves and cones or flowers. In flowering plants the gametophytes are very reduced in size, and are represented by the germinated pollen and the embryo sac.
Hornworts are a group of bryophytes constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, the flattened, green plant body of a hornwort is the gametophyte plant.
An antheridium is a haploid structure or organ producing and containing male gametes. The plural form is antheridia, and a structure containing one or more antheridia is called an androecium. Androecium is also the collective term for the stamens of flowering plants.
A pteridophyte is a vascular plant that disperses spores. Because pteridophytes produce neither flowers nor seeds, they are sometimes referred to as "cryptogams", meaning that their means of reproduction is hidden. Ferns, horsetails, and lycophytes are all pteridophytes. However, they do not form a monophyletic group because ferns are more closely related to seed plants than to lycophytes. "Pteridophyta" is thus no longer a widely accepted taxon, but the term pteridophyte remains in common parlance, as do pteridology and pteridologist as a science and its practitioner, respectively. Ferns and lycophytes share a life cycle and are often collectively treated or studied, for example by the International Association of Pteridologists and the Pteridophyte Phylogeny Group.
Pteridium aquilinum, also known as eagle fern, is a species of fern occurring in temperate and subtropical regions in both hemispheres. The extreme lightness of its spores has led to its global distribution.
Monoicy is a sexual system in haploid plants where both sperm and eggs are produced on the same gametophyte, in contrast with dioicy, where gametophytes produce only sperm or eggs but never both. Both monoicous and dioicous gametophytes produce gametes in gametangia by mitosis rather than meiosis, so that sperm and eggs are genetically identical with their parent gametophyte.
A prothallus, or prothallium, is usually the gametophyte stage in the life of a fern or other pteridophyte. Occasionally the term is also used to describe the young gametophyte of a liverwort or peat moss as well.
Plant reproduction is the production of new offspring in plants, which can be accomplished by sexual or asexual reproduction. Sexual reproduction produces offspring by the fusion of gametes, resulting in offspring genetically different from the parent or parents. Asexual reproduction produces new individuals without the fusion of gametes, genetically identical to the parent plants and each other, except when mutations occur.
Environmental sex determination is the establishment of sex by a non-genetic cue, such as nutrient availability, experienced within a discrete period after fertilization. Environmental factors which often influence sex determination during development or sexual maturation include light intensity and photoperiod, temperature, nutrient availability, and pheromones emitted by surrounding plants or animals. This is in contrast to genotypic sex determination, which establishes sex at fertilization by genetic factors such as sex chromosomes. Under true environmental sex determination, once sex is determined, it is fixed and cannot be switched again. Environmental sex determination is different from some forms of sequential hermaphroditism in which the sex is determined flexibly after fertilization throughout the organism’s life.
Heterospory is the production of spores of two different sizes and sexes by the sporophytes of land plants. The smaller of these, the microspore, is male and the larger megaspore is female. Heterospory evolved during the Devonian period from isospory independently in several plant groups: the clubmosses, the arborescent horsetails, and progymnosperms. This occurred as part of the process of evolution of the timing of sex differentiation.
Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes (haploid) combines with another to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). Sexual reproduction is the most common life cycle in multicellular eukaryotes, such as animals, fungi and plants. Sexual reproduction does not occur in prokaryotes, but they have processes with similar effects such as bacterial conjugation, transformation and transduction, which may have been precursors to sexual reproduction in early eukaryotes.
The word gametophore, also known as gametangiophore, is composed of gametangium and "phore". In moss and fern (Archegoniata) the gametophore is the bearer of the sex organs (gametangia), the female archegonia and the male antheridia. If both the archegonia and antheridia are arranged at the same plant, they are called monoicous. If there are female and male plants they are called dioicous.
Ceratopteris richardii is a fern species belonging to the genus Ceratopteris, one of only two genera of the subfamily Parkerioideae of the family Pteridaceae. It is one of several genera of ferns adapted to an aquatic existence. C. richardii was previously regarded as being part of the species Ceratopteris thalictroides.
Polyphlebium venosum, the veined bristle-fern or bristle filmy fern, is a fern in the family Hymenophyllaceae. It is only found in wet forests, mainly growing as an epiphyte on the shady side of the soft tree fern, Dicksonia antartica. It also grows on logs, trunks of trees and rarely on trunks of Cyathea species or on wet rock-faces. It is found in the wetter parts of Eastern Australia and New Zealand. P. venosum has poor long-distance dispersal compared to other ferns due to its short lived spore. Notable features of Polyphlebium venosum include it being one cell layer thick, 5–15 cm in length, having many branching veins and a trumpet shaped indusium.
Dioicy is a sexual system where archegonia and antheridia are produced on separate gametophytes. It is one of the two main sexual systems in bryophytes. One biologist stated there is a good reason for the distinction between dioecy and dioicy. Others have stated that the term dioecy is meaningless for bryophytes. Nonetheless dioecy and dioicy are comparable in many respects.