Saturday, March 9, 2013

It's allometric, my dear Watson

ResearchBlogging.orgGiant and colossal squid have the largest eyes of any living animals. Eyes are expensive organs to build and maintain, which led some researchers to suggest that there must be a strong evolutionary advantage for large eyes in giant squid. Using a mathematical model they found that giant squid eyes were best suited for detecting large dimly lit objects. They argued that the only stimulus that was both large enough and important enough for giant and colossal squid to detect was the light produced by bioluminescent organisms disturbed by hunting sperm whales.

A giant squid, Architeuthis dux (top), and a colossal squid, Mesonychoteuthis hamiltoni (bottom), being hauled up from the depths (images from National Geographic here and here respectively).
When I wrote about the paper, one of the criticisms I had was that the authors had failed to consider allometric scaling. Although the authors made comparisons of eye size with fish and extinct marine reptiles of similar size, they had not looked at eye size in other squid. Giant and colossal squid are the largest of all squid and their eyes could simply be large because they scaled up with their body size. I did a very crude analysis by conducting a literature search for papers that reported both eye size and body size in squid. From that I concluded that eye size was not disproportionately large relative to body size in giant and colossal squid.

Now, fortunately, nobody needs to rely on my poor-man's analysis. Schmitz et al. have published in BMC Evolutionary Biology that examines data from 87 different squid species and concludes that when allometric scaling is taken into account eye size in giant and colossal squids is not exceptional. In fact, it's pretty much exactly what you would expect if you scaled up another squid species to the same size. Indeed, there were a couple of groups, such as the bobtail squid, that had larger eyes relative to body size than the giant squid.

A regression of eye diameter on mantle length for 87 species of squid. Points for individual measurements in giant (yellow) and colossal (red) squid are shown for comparison (taken from Schmitz et al. 2013).
Further, Schmitz et al. also argue that many of the parameter values used in the original study are inappropriate. The original study based all of their optical performance calculations on the largest recorded giant squid eye diameter of 27 centimeters. But, this is problematic because the optical ability of such a large eye is likely to apply mainly to very large old squid, who are likely to have already reproduced. Eyes that only provide an advantage late in life are unlikely to contribute much to individual fitness. The original paper also probably set the values for the density and amount of light emitted from bioluminescent organisms in the deep sea too high.

When Schmitz et al. used more realistic values in the model they found that there was no unique advantage of large eyes for detecting large luminous objects, such as foraging sperm whales. Pupil sizes ranging from 2 centimeters up to the 15 centimeters used in the original model performed roughly equally well at detecting point sources and large luminous objects. Moreover, as eye size increased there was a slightly greater advantage for detecting point sources of light rather than large luminous objects. Thus, with more realistic parameter values, the conclusions of the original paper are essentially reversed.

References

Schmitz, L., Motani, R., Oufiero, C., Martin, C., McGee, M., Gamarra, A., Lee, J., & Wainwright, P. (2013). Allometry indicates giant eyes of giant squid are not exceptional BMC Evolutionary Biology, 13 (1) DOI: 10.1186/1471-2148-13-45  

Nilsson, D., Warrant, E., Johnsen, S., Hanlon, R., & Shashar, N. (2012). A Unique Advantage for Giant Eyes in Giant Squid Current Biology, 22 (8), 683-688 DOI: 10.1016/j.cub.2012.02.031

1 comment:

  1. Great work! I love it when science bloggers turn out to have been right & the peer-reviewed literature catches up later :)

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