Anisogamy (noun) (also called heterogamy) refers to a form of sexual reproduction involving the union or fusion of two dissimilar gametes (differing either in size and/or form) — anisogamous, anisogamic, (adj.). The smaller gamete is considered to be male (sperm cell), whereas the larger gamete is regarded as female (egg cell).
There are several types of anisogamy. Both gametes may be flagellated and thus motile. Alternatively, neither of the gametes may be flagellated. This situation occurs for example in some algae and plants. In the red alga Polysiphonia, large non-motile egg cells are fertilized by small, non-motile spermatia. In flowering plants, the gametes are non-motile cells within gametophytes.
The form of heterogamy that occurs in animals, including humans, is oogamy. In oogamy, a large, non-motile egg cell (ovum) is fertilized by a small, motile sperm cell (spermatozoon). The large egg cell is optimized for longevity, whereas the small sperm cell 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.
Evolution of anisogamy
Anisogamy is the phenomenon of fertilization of small gametes (sperm) and big gametes (eggs). Gamete size difference is the fundamental difference between males and females. Anisogamy first evolved in multicellular haploid species after the differentiation of different mating types had already been established. However, in Ascomycetes, anisogamy evolved from isogamy before mating types.
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. 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.
The first formal, mathematical theory proposed to explain the evolution of anisogamy was based on gamete limitation: this model assumed that natural selection would lead to gamete sizes that result in the largest population-wide number of successful fertilizations. 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.
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. 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 are parasitizing the large gametes that are left with the task of providing most of the 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. 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. 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).
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. Some comparative empirical evidence for the gamete competition theories exists, although it is difficult to use this evidence to fully tease apart the competition and limitation theories because their testable predictions are similar. It has also been claimed that some of the organisms used in such comparative studies do not fit the theoretical assumptions well.
- Dusenbery, David B. (2009). "Chapter 20". Living at Micro Scale. Cambridge, Mass: Harvard University Press. ISBN 978-0-674-03116-6.
- Beukeboom, L. & Perrin, N. (2014). The Evolution of Sex Determination. Oxford University Press, p. 25 . Online resources, .
- Lessells C.M., Snook R.R., Hosken D.J. 2009 The evolutionary origin and maintenance of sperm: selection for a small, motile gamete mating type. In Sperm biology: An evolutionary perspective (eds. Birkhead T.R., Hosken D.J., Pitnick S.), pp. 43-67. London, Academic press.
- Lehtonen J., Parker G.A. 2014 Gamete competition, gamete limitation, and the evolution of the two sexes. Molecular Human Reproduction 20 (12): 1161-1168. doi:10.1093/molehr/gau068
- Kalmus H. 1932 Über den Erhaltungswert der phänotypischen (morphologischen) Anisogamie und die Entstehung der ersten Geschlechtsunterschiede. Biol Zentralbl 52, 716–736.
- Scudo F.M. 1967 Adaptive value of sexual dimorphism - I, anisogamy. Evolution 21(2), 285-291.
- Dusenbery D.B. 2000 Selection for high gamete encounter rates explains the success of male and female mating types. Journal of Theoretical Biology 202(1), 1-10
- Williams G.C. 1966 Adaptation and natural selection: a critique of some current evolutionary thoughts. New Jersey, Princeton
- Parker G.A., Baker R.R., Smith V.G.F. 1972 The origin and evolution of gamete dimorphism and the male-female phenomenon. Journal of Theoretical Biology 36(3), 529-553.
- Parker G.A. 1978 Selection on non-random fusion of gametes during evolution of anisogamy. Journal of Theoretical Biology 73(1), 1-28.
- Parker G.A. 1982 Why are there so many tiny sperm? Sperm competition and the maintenance of two sexes. Journal of Theoretical Biology 96(2), 281-294.
- Cox P.A., Sethian J.A. 1985 Gamete motion, search, and the evolution of anisogamy, oogamy, and chemotaxis. American Naturalist 125(1), 74-101.
- Iyer P., Roughgarden J. 2008 Gametic conflict versus contact in the evolution of anisogamy. Theoretical Population Biology 73(4), 461-472. (doi:10.1016/j.tpb.2008.02.002).
- Yang J.-N. 2010 Cooperation and the evolution of anisogamy. Journal of Theoretical Biology 264(1), 24-36. (doi:http://dx.doi.org/10.1016/j.jtbi.2010.01.019).
- Lehtonen J., Kokko H. 2011 Two roads to two sexes: unifying gamete competition and gamete limitation in a single model of anisogamy evolution. Behav Ecol Sociobiol 65(3), 445-459. (doi:10.1007/s00265-010-1116-8).
- Parker G.A., Lehtonen J. 2014 Gamete evolution and sperm numbers: sperm competition versus sperm limitation. Proceedings of the Royal Society B: Biological Sciences 281(1791). (doi:10.1098/rspb.2014.0836).
- Knowlton N. 1974 A note on the evolution of gamete dimorphism. Journal of theoretical Biology 46(1), 283-285.
- Parker G.A. 2011 The origin and maintenance of two sexes (anisogamy), and their gamete sizes by gamete competition. In The evolution of anisogamy (eds. Togashi T., Cox P.A.), pp. 17-74. Cambridge, Cambridge University Press.
- Randerson J.P., Hurst L.D. 2001 The uncertain evolution of the sexes. Trends in Ecology & Evolution 16(10), 571-579.