werewolf

Author notes

17

marine

on Nov. 29, 2007

The testicle (from Latin testis, meaning "witness",[1] plural testes) is the male generative gland in animals.

Male mammals have two testicles, which are often contained within an extension of the abdomen called the scrotum. In mammals with external testicles it is most common for one testis to hang lower than the other. It is estimated that in about 85% of men the lower hanging testicle is the left one. This is due to differences in the vascular anatomical structure on the right and left sides.

The basal condition for mammals is to have internal testicles. Only the Boreoeutherian land mammals, the large group of mammals that includes humans, have externalized testicles. Indeed their testicles function best at temperatures lower than their core body temperature. Their testes are located outside of the body, suspended by the spermatic cord within the scrotum. The testes of the non-boreotherian mammals such as the monotremes, armadillos, sloths, elephants remain within the abdomen.[3] There are also some Boreoeutherian mammals with internal testes, such as the rhinoceros.

Marine boreotherian mammals such as whales and dolphins, also have internal testes, but it has recently been shown (e.g., for dolphins) that they use elaborate vascular networks to provide the necessary temperature lowering for optimum function. As external testes would increase drag, many boreotherian aquatic mammals have internal testes which are kept cool by special circulatory systems that cool the arterial blood going to the testes by placing the arteries near veins bringing cooled venous blood from the skin.

There are several hypotheses why most boreotherian mammals have external testes which operate best at a temperature that is slightly less than the core body temperature, e.g. that it is stuck with enzymes evolved in a colder temperature due to external testes evolving for different reasons, that the lower temperature of the testes simply is more efficient for sperm production.

1) More efficient. The classic hypothesis is that cooler temperature of the testes allows for more efficient fertile spermatogenesis. In other words, there are no possible enzymes operating at normal core body temperature that are as efficient as the ones evolved, at least none appearing in our evolution so far.

The early mammals had lower body temperatures and thus their testes worked efficiently within their body. However it is argued that boreotherian mammals have higher body temperatures than the other mammals and had to develop external testicles to keep them cool. It is argued that those mammals with internal testicles, such as the monotremes, armadillos, sloths, elephants, and rhinoceroses, have a lower core body temperatures than those mammals with external testicles.

However, the question remains why birds despite having very high core body temperatures have internal testes and did not evolve external testes.[4] It was once theorized that birds used their air sacs to cool the testes internally, but later studies revealed that birds' testes are able to function at core body temperature.[4]

2) Irreversible adaptation to sperm competition. It has been suggested that the ancestor of the boreoeutherian mammals was a small mammal that required very large testes (perhaps rather like those of a hamster) for sperm competition and thus had to place its testes outside the body.[5] This led to enzymes involved in spermatogenesis, spermatogenic DNA polymerase beta and recombinase activities evolving a unique temperature optimum, slightly less than core body temperature. When the boreoeutherian mammals then diversified into forms that were larger and/or did not require intense sperm competition they were stuck with enzymes that operated best at cooler temperatures and had to keep their testicles outside the body.

3) Protection from abdominal cavity pressure changes. One argument for the evolution of external testes is that it protects the testes from abdominal cavity pressure changes caused by jumping and galloping.