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How to Design Experiments in Animal Behaviour∗ 12

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How to Design Experiments in Animal Behaviour∗ 12 SERIES ARTICLE How to Design Experiments in Animal Behaviour∗ 12. Male Frogs Sing for Sex but Why Don’t They Do Their Best? Raghavendra Gadagkar Charles Darwin proposed a separate theory of sexual selec- tion, as distinct from his theory of natural selection, to ac- count for adaptations that confer success in finding a mate, which may sometimes be quite the opposite of what is best for survival. Darwin’s proposal that females have a sense of beauty and choose mating partners that appear beautiful to them was met with much scepticism. But today we have a rather detailed understanding of what animals find beauti- ful and why. In this article, I will describe a few very sim- Raghavendra Gadagkar is ple experiments performed by Michael J. Ryan, in collabora- DST Year of Science Chair Professor at the Centre for tion with A. Stanley Rand, herpetologist extraordinaire and Ecological Sciences, Indian Merlin D Tuttle of the Bat Conservation International fame, Institute of Science, that laid the foundation for our current understanding of the Bangalore, Honorary meaning and evolution of beauty. Studying the tungara´ frog Professor at JNCASR, and Non-Resident Permanent on Barro Colorado Island, a research station of the Smithso- Fellow of the nian Tropical Research Institute in Panama, they showed that Wissenschaftskolleg (Institute (1) male tungara´ frogs can produce both simple calls, consist- for Advanced Study), Berlin. ing of just a whine, or complex calls in which one or more During the past 40 years he has established an active chucks are added to the whine, (2) female tungara´ frogs have school of research in the area a decided preference to mate with males giving complex calls, of animal behaviour, ecology (3) males are nevertheless reluctant to add chucks to their and evolution. The origin and calls and generally do so only when they hear other males evolution of cooperation in animals, especially in social calling, and (4) the local predatory fringe-lipped bat also has insects, such as ants, bees and a decided preference to eat males giving complex calls. Male wasps, is a major goal of his tungara´ frogs thus face a trade-off between sex and survival. research. // / / These experiments not only answered the question of why http: ces.iisc.ac.in hpg ragh. https://www.researchgate.net/ males don’t do their best when it comes to singing, but they profile/Raghavendra Gadagkar also set the stage for many more sophisticated investigations ∗Vol.25, No.6, DOI: https://doi.org/10.1007/s12045-020-0998-7 RESONANCE | June 2020 817 SERIES ARTICLE that have led to an understanding of how and why natural Keywords selection has favoured this particular sexual aesthetic in the T´ungara frogs, mating call, frogs and this particular culinary aesthetic in the bats. Physalaemus pustulosus, sexual selection, female choice. The Cost of Sex Nearly all multicellular organisms practice sexual reproduction— the union of male and female haploid gametes to form a diploid zygote in every generation. Very few multicellular species rou- tinely use the ancient practice of asexual reproduction, although many have retained it for use under certain conditions. This is something of an evolutionary paradox because sexual reproduc- tion is costly—it requires two parents to produce offspring. Un- like in most sexual species where about half the population con- sists of males, in an asexual population all individuals are iden- tical (not differentiated into males and females, but traditionally referred to as females because they give birth to offspring) and reproduce all by themselves. This two-fold cost of sex is the sub- ject of much research and many theories. That apart, there is also the problem in sexually reproducing species that males and fe- males have to find each other in every single generation. Every time I see an obscure insect I can’t help marvelling at the fact that at least some members of that species must have found sex- It is a curious fact of the ual partners in every generation without fail, for the millions of biology of sexual years that the species has been around. Not surprisingly, sexually reproduction that the reproducing organisms go to a great deal of trouble to ensure the burden of finding a sexual partner falls more union of males and females. heavily on males than on It is a curious fact of the biology of sexual reproduction that the females. Because females generally burden of finding a sexual partner falls more heavily on males produce relatively small than on females. Because females generally produce relatively numbers of costly eggs small numbers of costly eggs and devote relatively more time and devote relatively and effort in caring for their offspring, they are the limiting re- more time and effort in caring for their offspring, source. Males on the other hand, generally produce large num- they are the limiting bers of cheap sperm, spend relatively less time and effort caring resource. for their offspring and can, therefore, mate with many more part- ners than is possible for a typical female. The mass of a nutrient- 818 RESONANCE | June 2020 SERIES ARTICLE filled human egg, for instance, is about a million times the mass of a human sperm, which contains little more than a haploid set of chromosomes! In this asymmetric situation, success in finding a sexual partner is relatively more uniformly distributed among the females but is highly skewed among the males—while most fe- males will find a sexual partner, some males will find many part- ners and many males will inevitably be left without any. Males, therefore, have to work much harder to find a mate—they either have to fight with other males in order to win in the male-male competition and/or they have to advertise themselves to the fe- males and emerge successfully through the filter of female choice. Darwin realised that what it takes to succeed in the sexual market is not always the same that is required to succeed in the struggle for survival. He, therefore, proposed a separate theory of Sexual Selection, to account for adaptations that confer success in find- ing a mate, which may sometimes be quite the opposite of what is best for survival. Perhaps not so surprisingly, male-male competition for females has been much better studied while female choice has been rather neglected, at least until recently. Darwin’s proposal that females have a sense of beauty and choose mating partners that appear beautiful to them was met with much skepticism, including by Alfred Russel Wallace, the co-discoverer of the principle of natu- ral selection. That females can indeed choose, and have a sense of beauty is the theme of this article, but of course, we will mainly use it as a context in which to continue to explore the practice of science and illustrate more ways of designing experiments and doing low-cost research. In particular, we will dwell on how male frogs advertise their beauty and what female frogs find beautiful. Frogs may seem an unlikely subject to explore beauty. Our com- mon perception of frogs is perhaps not one they would be proud of! There are over 7000 species of frogs, and a great deal is There are over 7000 known about them—at least biologists have found them worthy species of frogs, and a of attention. In 2007 Professor Kentwood D Wells of the Uni- great deal is known about them—at least versity of Connecticut in the USA published a 1400-page book biologists have found entitled The Ecology & Behaviour of Amphibians [1], which has them worthy of attention. RESONANCE | June 2020 819 SERIES ARTICLE Figure 1. Michael J Ryan (1953–). Born to a truck driver in Bronx, New York as the oldest of eleven sib- lings, Michael J Ryan rose to become the Clark Hubbs Re- gents Professor in Zoology at the University of Texas, Austin, USA. Michael Ryan has spent a lifetime study- ing the t´ungara frog in an attempt to understand what is beauty and how and why some traits come to be re- been described as “the definitive one-volume work on the world’s garded as beautiful while amphibians”. Professor Wells begins his book with an amazing others not. His research, quote by Carl von Linnaeus, the father of taxonomy: “These foul culminating in the theory and loathsome animals ... are abhorrent because of their cold of sensory exploitation, has body, pale colour, cartilaginous skeleton, filthy skin, fierce as- shown that while males have pect, calculating eye, offensive smell, harsh voice, squalid habita- exploited the sensory bi- tion, and terrible venom; and so their creator has not exerted His ases of females over evolu- powers to make many of them.” tionary time, at the present time females are the bi- This reputation of amphibians apparently did not deter Professor ological puppeteers, mak- Michael J. Ryan (Figure 1), one of the protagonists of this article, ing the males sing exactly to choose a frog to explore beauty. Clearly, he made a wise choice what their brains desire—a as his 40-year research [2] on the T´ungara Frog has led recently to paradigm shift in the study the publication of his fascinating book, A Taste for the Beautiful: of sexual selection. The lat- The Evolution of Attraction [3]. ter part of A Taste for the Beautiful: The Evolution of Attraction [3] was written The Tungara´ Frog after Mike suffered a serious accident which has left him Physalaemus pustulosus is a small, brownish frog, about 3 cm in a wheelchair with a spinal in length (length of frogs is usually measured as the distance be- cord injury.
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