Anisogamy

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Different forms of anisogamy: A) anisogamy of motile cells, B) oogamy (egg cell and sperm cell), C) anisogamy of non-motile cells (egg cell and spermatia). Anisogamy.svg
Different forms of anisogamy: A) anisogamy of motile cells, B) oogamy (egg cell and sperm cell), C) anisogamy of non-motile cells (egg cell and spermatia).

Anisogamy is a form of sexual reproduction that involves the union or fusion of two gametes that differ in size and/or form. The smaller gamete is male, a sperm cell, whereas the larger gamete is female, typically an egg cell. Anisogamy is predominant among multicellular organisms. [1] In both plants and animals, gamete size difference is the fundamental difference between females and males. [2]

Contents

Anisogamy most likely evolved from isogamy. [3] Since the biological definition of male and female is based on gamete size, the evolution of anisogamy is viewed as the evolutionary origin of male and female sexes. [4] [5] Anisogamy is an outcome of both natural selection and sexual selection, [6] and led the sexes to different primary and secondary sex characteristics [7] including sex differences in behavior. [8]

Geoff Parker, Robin Baker, and Vic Smith were the first to provide a mathematical model for the evolution of anisogamy that was consistent with modern evolutionary theory. [4] Their theory was widely accepted but there are alternative hypotheses about the evolution of anisogamy. [9] [1]

Etymology

Anisogamy (the opposite of isogamy ) comes from the ancient Greek negative prefix a(n)- (alpha privative), the Greek adjective isos (meaning equal) and the Greek verb gameo (meaning to have sex/to reproduce), eventually meaning "non-equal reproduction" obviously referring to the enormous differences between male and female gametes in size and abilities. [10] The first known use of the term anisogamy was in the year 1891.[ clarification needed ] [11]

Definition

Anisogamy is the form of sexual reproduction that involves the union or fusion of two gametes which differ in size and/or form. [12] The smaller gamete is considered to be male (a sperm cell), whereas the larger gamete is regarded as female (typically an egg cell, if non-motile). [13] [14]

There are several types of anisogamy. Both gametes may be flagellated and therefore motile. Alternatively, as in flowering plants, conifers and gnetophytes, neither of the gametes are flagellated. In these groups, the male gametes are non-motile cells within pollen grains, and are delivered to the egg cells by means of pollen tubes. In the red alga Polysiphonia , non-motile eggs are fertilized by non-motile sperm.

The form of anisogamy that occurs in animals, including humans, is oogamy, where a large, non-motile egg (ovum) is fertilized by a small, motile sperm (spermatozoon). The egg is optimized for longevity, whereas the small sperm is optimized for motility and speed. The size and resources of the egg cell allow for the production of pheromones, which attract the swimming sperm cells. [15]

Sexual dimorphism

Anisogamy is a core element of sexual dimorphism that helps to explain phenotypic differences between sexes. [16] [17] Researchers estimate that over 99.99% of eukaryotes reproduce sexually. [18] Most do so by way of male and female sexes, both of which are optimized for reproductive potential. Due to their differently sized and shaped gametes, both males and females have developed physiological and behavioral differences that optimize the individual's fecundity. [16] Since most egg laying females typically must bear the offspring and have a more limited reproductive cycle, this typically makes females a limiting factor in the reproductive success rate of males in a species. This process is also true for females selecting males, and assuming that males and females are selecting for different traits in partners, would result in phenotypic differences between the sexes over many generations. This hypothesis, known as the Bateman's Principle, is used to understand the evolutionary pressures put on males and females due to anisogamy. [19] Although this assumption has criticism, it is a generally accepted model for sexual selection within anisogamous species. The selection for different traits depending on sex within the same species is known as sex-specific selection, and accounts for the differing phenotypes found between the sexes of the same species. This sex-specific selection between sexes over time also leads to the development of secondary sex characteristics, which assist males and females in reproductive success.

In most species, both sexes choose mates based on the available phenotypes of potential mates. [19] These phenotypes are species-specific, resulting in varying strategies for successful sexual reproduction. For example, large males are sexually selected for in elephant seals because their large size helps the male fight off other males, but small males are sexually selected for in spiders for they can mate with the female more quickly while avoiding sexual cannibalism. [20] However, despite the large range of sexually selected phenotypes, most anisogamous species follow a set of predictable desirable traits and selective behaviors based on general reproductive success models.

Female phenotypes

For internal fertilizers, female investment is high in reproduction since they typically expend more energy throughout a single reproductive event. This can be seen as early as oogenesis, for the female sacrifices gamete number for gamete size to better increase the survival chances of the potential zygote; a process more energetically demanding than spermatogenesis in males. [21] Oogenesis occurs in the ovary, a female-specific organ that also produces hormones to prepare other female-specific organs for the changes necessary in the reproductive organs to facilitate egg delivery in external fertilizers, and zygote development in internal fertilizers. The egg cell produced is not only large, but sometimes even immobile, requiring contact with the more mobile sperm to instigate fertilization. [21]

Since this process is very energy-demanding and time-consuming for the female, mate choice is often integrated into the female's behavior. [16] Females will often be very selective of the males they choose to reproduce with, for the phenotype of the male can be indicative of the male's physical health and heritable traits. Females employ mate choice to pressure males into displaying their desirable traits to females through courtship, and if successful, the male gets to reproduce. This encourages males and females of specific species to invest in courtship behaviors as well as traits that can display physical health to a potential mate. This process, known as sexual selection, [16] results in the development of traits to ease reproductive success rather than individual survival, such as the inflated size of a termite queen. It is also important for females to select against potential mates that may have a sexually transmitted infection, for the disease could not only hurt the female's reproductive ability, but also damage the resulting offspring. [22]

Although not uncommon in males, females are more associated with parental care. Since females are on a more limited reproductive schedule than males, a female often invests more in protecting the offspring to sexual maturity than the male. Like mate choice, the level of parental care varies greatly between species, and is often dependent on the number of offspring produced per sexual encounter. [23]

In many species, including ones from all major vertebrate groups [24] females can utilize sperm storage, [25] a process by which the female can store excess sperm from a mate, and fertilize her eggs long after the reproductive event if mating opportunities drop or quality of mates decreases. By being able to save sperm from more desirable mates, the female gains more control over its own reproductive success, thus allowing for the female to be more selective of males as well as making the timing of fertilization potentially more frequent if males are scarce. [25]

Male phenotypes

For males of all species, the sperm cells they produce are optimized for ensuring fertilization of the female egg. These sperm cells are created through spermatogenesis, a form of gametogenesis that focuses on developing the most possible gametes per sexual encounter. [21] Spermatogenesis occurs in the testis, a male specific organ that also produces hormones that trigger the development of secondary sex characteristics. Since the male's gametes are energetically cheap and abundant in every ejaculation, a male can greatly increase his sexual success by mating far more frequently than the female. [21] Sperm, unlike egg cells, are also mobile, allowing for the sperm to swim towards the egg through the female's sexual organs. Sperm competition is also a major factor in the development of sperm cells. Only one sperm can fertilize an egg, and since females can potentially mate with more than one male before fertilization occurs, producing sperm cells that are faster, more abundant, and more viable than that produced by other males can give a male reproductive advantage. [21]

Since females are often the limiting factor in a species reproductive success, males are often expected by the females to search and compete for the female, known as intraspecific competition. [19] This can be seen in organisms such as bean beetles, as the male that searches for females more frequently is often more successful at finding mates and reproducing. In species undergoing this form of selection, a fit male would be one that is fast, has more refined sensory organs, and spatial awareness. [19]

Some secondary sex characteristics are not only meant for attracting mates, but also for competing with other males for copulation opportunities. Some structures, such as antlers in deer, can provide benefits to the male's reproductive success by providing a weapon to prevent rival males from achieving reproductive success. [22] However, other structures such as the large colorful tail feathers found in male peacocks, are a result of Fisherian runaway as well as several more species specific factors. Due to females selecting for specific traits in males, over time, these traits are exaggerated to the point where they could hinder the male's survivability. [22] However, since these traits greatly benefit sexual selection, their usefulness in providing more mating opportunities overrides the possibility that the trait could lead to a shortening of its lifespan through predation or starvation. These desirable traits extend beyond physical body parts, and often extend into courtship behavior and nuptial gifts as well.

Although some behaviors in males are meant to work within the parameters of female choice, some male traits work against it. Strong enough males, in some cases, can force themselves upon a female, forcing fertilization and overriding female choice. [26] Since this can often be dangerous for the female, an evolutionary arms race between the sexes is often an outcome.

History

Charles Darwin wrote that anisogamy had an impact on the evolution of sexual dimorphism. He also argued that anisogamy had an impact on sexual behavior. [27] Anisogamy first became a major topic in the biological sciences when Charles Darwin wrote about sexual selection.[ clarification needed ] [28]

Mathematical models seeking to account for the evolution of anisogamy were published as early as 1932, but the first model consistent with evolutionary theory was that published by Geoff Parker, Robin Baker and Vic Smith in 1972. [4]

Evolution

Although its evolution has left no fossil records, [3] it is generally accepted that anisogamy evolved from isogamy and that it has evolved independently in several groups of eukaryotes including protists, algae, plants and animals. [29] According to John Avise anisogamy probably originated around the same time sexual reproduction and multicellularity occurred, [30] over 1 billion years ago. [31] Anisogamy first evolved in multicellular haploid species after different mating types had become established. [32]

The three main theories for the evolution of anisogamy are gamete competition, gamete limitation, and intracellular conflicts, but the last of these three is not well supported by current evidence. [33] Both gamete competition and gamete limitation assume that anisogamy originated through disruptive selection acting on an ancestral isogamous population with external fertilization, due to a trade-off between larger gamete number and gamete size (which in turn affects zygote survival), because the total resource one individual can invest in reproduction is assumed to be fixed. [31]

The first formal, mathematical theory proposed to explain the evolution of anisogamy was based on gamete limitation: [34] this model assumed that natural selection would lead to gamete sizes that result in the largest population-wide number of successful fertilizations. [34] [35] [36] If it is assumed that a certain amount of resources provided by the gametes are needed for the survival of the resulting zygote, and that there is a trade-off between the size and number of gametes, then this optimum was shown to be one where both small (male) and large (female) gametes are produced. However, these early models assume that natural selection acts mainly at the population level, something that is today known to be a very problematic assumption. [37]

The first mathematical model to explain the evolution of anisogamy via individual level selection, and one that became widely accepted was the theory of gamete or sperm competition. [38] [39] [40] Here, selection happens at the individual level: those individuals that produce more (but smaller) gametes also gain a larger proportion of fertilizations simply because they produce a larger number of gametes that 'seek out' those of the larger type. However, because zygotes formed from larger gametes have better survival prospects, this process can again lead to the divergence of gametes sizes into large and small (female and male) gametes. The end result is one where it seems that the numerous, small gametes compete for the large gametes that are tasked with providing maximal resources for the offspring.

Some recent theoretical work has challenged the gamete competition theory, by showing that gamete limitation by itself can lead to the divergence of gamete sizes even under selection at the individual level. [41] [42] [43] While this is possible, it has also been shown that gamete competition and gamete limitation are the ends of a continuum of selective pressures, and they can act separately or together depending on the conditions. [44] These selection pressures also act in the same direction (to increase gamete numbers at the expense of size) and at the same level (individual selection). Theory also suggests that gamete limitation could only have been the dominant force of selection for the evolutionary origin of the sexes under quite limited circumstances, and the presence on average of just one competitor can makes the 'selfish' evolutionary force of gamete competition stronger than the 'cooperative' force of gamete limitation even if gamete limitation is very acute (approaching 100% of eggs remaining unfertilized). [45]

There is then a relatively sound theory base for understanding this fundamental transition from isogamy to anisogamy in the evolution of reproduction, which is predicted to be associated with the transition to multicellularity. In fact, Hanschen et al. (2018) demonstrate that anisogamy evolved from isogamous multicellular ancestors and that anisogamy would subsequently drive secondary sexual dimorphism. [32] Some comparative empirical evidence for the gamete competition theories exists, [33] [46] [47] although it is difficult to use this evidence to fully tease apart the competition and limitation theories because their testable predictions are similar. [31] It has also been claimed that some of the organisms used in such comparative studies do not fit the theoretical assumptions well. [48]

A valuable model system to the study of the evolution of anisogamy is the volvocine algae, which group of chlorophytes is quite unique for its extant species exhibit a diversity of mating systems (isogamy and anisogamy) in addition to its extremes in both unicellularity and multicellularity with a diversity of forms in species of intermediate ranges of sizes. [49] Marine algae have been closely studied to understand the trajectories of such diversified reproductive systems, [50] evolution of sex and mating types, [51] as well as the adaptiveness and stability of anisogamy. [52] [50] [32]

See also

Related Research Articles

<span class="mw-page-title-main">Asexual reproduction</span> Reproduction without a sexual process

Asexual reproduction is a type of reproduction that does not involve the fusion of gametes or change in the number of chromosomes. The offspring that arise by asexual reproduction from either unicellular or multicellular organisms inherit the full set of genes of their single parent and thus the newly created individual is genetically and physically similar to the parent or an exact clone of the parent. Asexual reproduction is the primary form of reproduction for single-celled organisms such as archaea and bacteria. Many eukaryotic organisms including plants, animals, and fungi can also reproduce asexually. In vertebrates, the most common form of asexual reproduction is parthenogenesis, which is typically used as an alternative to sexual reproduction in times when reproductive opportunities are limited. Some monitor lizards, including Komodo dragons, can reproduce asexually.

<span class="mw-page-title-main">Gamete</span> Haploid sex cell

A gamete 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. The name gamete was introduced by the German cytologist Eduard Strasburger in 1878.

<span class="mw-page-title-main">Gametophyte</span> Haploid stage in the life cycle of plants and algae

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.

<span class="mw-page-title-main">Reproduction</span> Biological process by which new organisms are generated from one or more parent organisms

Reproduction is the biological process by which new individual organisms – "offspring" – are produced from their "parent" or parents. There are two forms of reproduction: asexual and sexual.

<span class="mw-page-title-main">Sex</span> Trait that determines an organisms sexually reproductive function

Sex is the biological trait that determines whether a sexually reproducing organism produces male or female gametes. During sexual reproduction, a male and a female gamete fuse to form a zygote, which develops into an offspring that inherits traits from each parent. By convention, organisms that produce smaller, more mobile gametes are called male, while organisms that produce larger, non-mobile gametes are called female. An organism that produces both types of gamete is hermaphrodite.

<span class="mw-page-title-main">Sexual dimorphism</span> Evolved difference in sex-specific characteristics

Sexual dimorphism is the condition where sexes of the same species exhibit different morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecious species, which consist of most animals and some plants. Differences may include secondary sex characteristics, size, weight, color, markings, or behavioral or cognitive traits. Male-male reproductive competition has evolved a diverse array of sexually dimorphic traits. Aggressive utility traits such as "battle" teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to sexual selection and natural selection. The opposite of dimorphism is monomorphism, when both biological sexes are phenotypically indistinguishable from each other.

<span class="mw-page-title-main">Sperm competition</span> Reproductive process

Sperm competition is the competitive process between spermatozoa of two or more different males to fertilize the same egg during sexual reproduction. Competition can occur when females have multiple potential mating partners. Greater choice and variety of mates increases a female's chance to produce more viable offspring. However, multiple mates for a female means each individual male has decreased chances of producing offspring. Sperm competition is an evolutionary pressure on males, and has led to the development of adaptations to increase male's chance of reproductive success. Sperm competition results in a sexual conflict between males and females. Males have evolved several defensive tactics including: mate-guarding, mating plugs, and releasing toxic seminal substances to reduce female re-mating tendencies to cope with sperm competition. Offensive tactics of sperm competition involve direct interference by one male on the reproductive success of another male, for instance by mate guarding or by physically removing another male's sperm prior to mating with a female. For an example, see Gryllus bimaculatus.

<span class="mw-page-title-main">Isogamy</span> Sexual reproduction form involving gametes of the same size

Isogamy is a form of sexual reproduction that involves gametes of the same morphology, and is found in most unicellular eukaryotes. Because both gametes look alike, they generally cannot be classified as male or female. Instead, organisms that reproduce through isogamy are said to have different mating types, most commonly noted as "+" and "−" strains.

<span class="mw-page-title-main">Male</span> Sex of an organism which produces sperm

Male is the sex of an organism that produces the gamete known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilisation. A male organism cannot reproduce sexually without access to at least one ovum from a female, but some organisms can reproduce both sexually and asexually. Most male mammals, including male humans, have a Y chromosome, which codes for the production of larger amounts of testosterone to develop male reproductive organs.

<span class="mw-page-title-main">Fish reproduction</span> Reproductive physiology of fishes

Fish reproductive organs include testes and ovaries. In most species, gonads are paired organs of similar size, which can be partially or totally fused. There may also be a range of secondary organs that increase reproductive fitness. The genital papilla is a small, fleshy tube behind the anus in some fishes, from which the sperm or eggs are released; the sex of a fish can often be determined by the shape of its papilla.

<span class="mw-page-title-main">Oogamy</span> Form of sexual reproduction

Oogamy is a form of anisogamy where the gametes differ in both size and form. In oogamy the large female gamete is immotile, while the small male gamete is mobile. Oogamy is a common form of anisogamy, with almost all animals and land plants being oogamous.

Monogamous pairing in animals refers to the natural history of mating systems in which species pair bond to raise offspring. This is associated, usually implicitly, with sexual monogamy.

<span class="mw-page-title-main">Sexual conflict</span> Term in evolutionary biology

Sexual conflict or sexual antagonism occurs when the two sexes have conflicting optimal fitness strategies concerning reproduction, particularly over the mode and frequency of mating, potentially leading to an evolutionary arms race between males and females. In one example, males may benefit from multiple matings, while multiple matings may harm or endanger females due to the anatomical differences of that species. Sexual conflict underlies the evolutionary distinction between male and female.

<span class="mw-page-title-main">Female</span> Sex of an organism that produces ova

An organism's sex is female if it produces the ovum, the type of gamete that fuses with the male gamete during sexual reproduction.

<span class="mw-page-title-main">Sexual reproduction</span> Biological process

Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete with a single set of chromosomes combines with another gamete to produce a zygote that develops into an organism composed of cells with two sets of chromosomes (diploid). This is typical in animals, though the number of chromosome sets and how that number changes in sexual reproduction varies, especially among plants, fungi, and other eukaryotes.

<span class="mw-page-title-main">Female sperm storage</span>

Female sperm storage is a biological process and often a type of sexual selection in which sperm cells transferred to a female during mating are temporarily retained within a specific part of the reproductive tract before the oocyte, or egg, is fertilized. This process takes place in some species of animals. The site of storage is variable among different animal taxa and ranges from structures that appear to function solely for sperm retention, such as insect spermatheca and bird sperm storage tubules, to more general regions of the reproductive tract enriched with receptors to which sperm associate before fertilization, such as the caudal portion of the cow oviduct containing sperm-associating annexins. Female sperm storage is an integral stage in the reproductive process for many animals with internal fertilization. It has several documented biological functions including:

Sexual antagonistic co-evolution is the relationship between males and females where sexual morphology changes over time to counteract the opposite's sex traits to achieve the maximum reproductive success. This has been compared to an arms race between sexes. In many cases, male mating behavior is detrimental to the female's fitness. For example, when insects reproduce by means of traumatic insemination, it is very disadvantageous to the female's health. During mating, males will try to inseminate as many females as possible, however, the more times a female's abdomen is punctured, the less likely she is to survive. Females that possess traits to avoid multiple matings will be more likely to survive, resulting in a change in morphology. In males, genitalia is relatively simple and more likely to vary among generations compared to female genitalia. This results in a new trait that females have to avoid in order to survive.

Interlocus sexual conflict is a type of sexual conflict that occurs through the interaction of a set of antagonistic alleles at two or more different loci, or the location of a gene on a chromosome, in males and females, resulting in the deviation of either or both sexes from the fitness optima for the traits. A co-evolutionary arms race is established between the sexes in which either sex evolves a set of antagonistic adaptations that is detrimental to the fitness of the other sex. The potential for reproductive success in one organism is strengthened while the fitness of the opposite sex is weakened. Interlocus sexual conflict can arise due to aspects of male–female interactions such as mating frequency, fertilization, relative parental effort, female remating behavior, and female reproductive rate.

<span class="mw-page-title-main">Social selection</span> Term used in biology

Social selection is a term used with varying meanings in biology.

Gynogenesis, a form of parthenogenesis, is a system of asexual reproduction that requires the presence of sperm without the actual contribution of its DNA for completion. The paternal DNA dissolves or is destroyed before it can fuse with the egg. The egg cell of the organism is able to develop, unfertilized, into an adult using only maternal genetic material. Gynogenesis is often termed "sperm parasitism" in reference to the somewhat pointless role of male gametes. Gynogenetic species, "gynogens" for short, are unisexual, meaning they must mate with males from a closely related bisexual species that normally reproduces sexually.

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