Gender origin

Definitions

For what the term 'gender' means in this essay, see the 'gender meaning' essay.

Gender origin

There are a number of theories about the origin of gender:

One theory suggests males differ from females primarily as a result of disruptive selection acting on gamete size.
This theory originated with Parker, Baker, and Smith in 1972 [1].
Their theory suggests males differ from females primarily according to whether they have small or large gametes.
It suggests isogamy (having gametes of equal size) is rare because it is not a stable strategy.
Disruptive selection causes large nutrient- containing gametes to evolve in conjunction with small nutrient-less exploiters.
Another theory proposes that gamete encounter rates explain the difference in size - rather than nutrient contributions.
This theory seems to be due to D Dusenbery [2, 3].
Another theory holds that males differ from females as a result of selection against pathogens favouring a single cell bottleneck.
Reproduction via cell fusion has the potential to allow pathogens from both parents into the offspring - increasing its risk of disease.
An example of vertical pathogen transmission through egg cytoplasm in mammals is the lymphocytic choriomeningitis virus.
Strip searching one set of genes minimizes the chances of this happening.
Another theory involves conflict between nuclear and cytoplasmic genes.
This theory is associated with Leda Cosmides and John Tooby [4].
Their theory suggests that males differ from females primarily according to whether their offspring inherit their organelles or not.
Lastly, there is the theory that gender exists to prevent inbreeding and selfing.

Prevent inbreeding

Gender exists to help prevent inbreeding - in many fungi.

These use 'mating types' to allow organisms to avoid mating with themselves and their near relatives.

The genes responsible often involve multiple loci on different chromosomes - and many alleles.

The result is a system which allows organisms to breed with most of the rest of the population, except for all but a small number of relatives.

While this indicates that gender exists to help prevent inbreeding in fungi, it also indicates strongly that it does not play that role very effectively in most animals and plants.

Having only two genders is a hopelessly inefficient system if preventing inbreeding is the aim.

Under such a scheme, selfing is prevented - but you can still breed with half of your closest relatives, and mate choice in the general population is needlessly restricted.

A system with multiple mating types works much better if preventing inbreeding is the goal. The two genders of most plants and animals must exist for some other reason.

Contingency vs inevitability

In the case of the cytoplasmic genes theory, gender is a consequence of particular contingent events during the early history of life, that resulted in living organisms forming a symbiotic union.

The disruptive selection theory instead suggests that gender is practically inevitable for complex organisms.

So: which is it, contingency or inevitability?

Inevitability gets my vote.

Even if organisms had no mitochondria, gender would still exist, due to all the adaptive reasons favouring it, as detailed in the list of theories of the origin of anisogamy, above.

From this perspective, mitochondrial conflict is regarded as being more likely to be to do with how gender exists in living organisms than why it exists - since it would still exist even if mitochondria did not.

Even our distant descendants - or advanced alien species - will still make extensive use systems where there is a division of labour between direct production of offspring, and propagation of information about how best to construct those offspring.

It is that division of labour which best characterises male and female roles - not anything to do with organelles.

Conventional view

I think most biologists regard gamete size as the most fundamental difference between males and females.

Richard Dawkins expresses this point of view in The Selfish Gene:

One group of individuals has large sex cells, and it is convenient to use the term female for them. The other group, which is convenient to call male has small sex cells. The difference is especially pronounced in reptiles and birds, where a single egg cell is big enough and nutritious enough to feed a developing embryo for several weeks. Even in humans, where the egg is microscopic, it is still many times larger than the sperm. As we shall see, it is possible to regard all the other differences between the sexes as stemming from this one basic difference. - Richard Dawkins, The Selfish Gene, 2nd ed. page 142.[5]

He goes on to cite: "The evolution of anisogamy: a game-theoretic approach" [6] - a paper about the disruptive selection theory.

Gene conflict theory popularity

I note that many modern writers seem to have embraced Leda Cosmides and John Tooby's more recent theory as though it is the whole truth.

In The Red Queen [7] Matt Ridley asks:

To ask 'Why are there two genders?' is to ask 'Why are mitochondria inherited through the maternal line.'

Such a statement assumes the Cosmides/Tooby perspective - that gender is all down to organelles.

He goes on to write:

Two genders have been invented, killer which provides the organelles, and victim which does not.

Similarly, in Mendel's Demon [8], Mark Ridley says:

The deep nature of maleness is to eject organelles from reproductive cells; The deep nature of femaleness is to keep them.

He goes on to make it clear that he regards gender as a contingency:

The theory implies that the existence of gender itself, in life on Earth, is an evolutionary fluke.

He says it depends on two accidents:

To be more exact, gender depends on two flukes, or events that look like flukes. One is that complex life evolved via the merger; the second is that both DNA sets of the merger partners were retained in the post-merger cell [...]

Conclusion

The comments above by modern authors ignore the adaptive reasons for anisogamy and the existence of gender - and are therefore seriously in error.

The story about organelles is an implementation detail - not the sole cause.

Gender and anisogamy would still exist if organelles did not.

References

Tim Tyler | Contact | http://alife.co.uk/