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Nikolaas Tinbergen the Careful Scientist

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Nikolaas Tinbergen the Careful Scientist GENERAL ARTICLE Nikolaas Tinbergen The Careful Scientist Sindhu Radhakrishna What is it about the work of some great scientists that sets them apart from the admirable work done by many good sci- entists? Is it because some, if not all, of their contributions stand the test of time? Or is it because their work paved the way for other great ideas to develop? Nikolaas Tinbergen’s life and work suggests that it is a combination of both these factors that characterise a truly great scientist. Nikolaas Tinbergen (1907–1988) was a Dutch scientist who stud- ied animal behaviour. Tinbergen began his academic career at Sindhu Radhakrishna holds a master’s degree in psychology the University of Leiden in the Netherlands and later moved to and a doctorate in animal Oxford, where he set up a school for animal behaviour stud- behaviour. She is presently ies. Tinbergen pioneered and popularised the science of ethol- Professor and Dean at the ogy through his research and teaching. He lectured widely on Animal Behaviour and Cognition Programme, the subject and wrote extensively; some of his more well-known National Institute of books include The Study of Instinct (1951), The Herring Gull’s Advanced Studies, World (1953), Curious Naturalists (1958), and the Time-life vol- Bengaluru. Her research ume, Animal Behavior (1965). Tinbergen was an influential bi- interests are in the fields of primatology, behavioral ologist within his lifetime and received many honours for his ecology and conservation work, most notably, the Nobel Prize, the Swammerdam Medal, biology, and her work is the Distinguished Scientific Contribution Award from the Ameri- focused on gaining a better can Psychological Association and the Diploma of Honour of the understanding of social behaviour and Sociedad Argentina Protectora de Animales. Apart from being communication in nocturnal an excellent communicator who mentored a new generation of primates. renowned biologists, Tinbergen was also an exceptional photog- rapher and filmmaker who was celebrated for his films on animal behaviour. Tinbergen is often described as one of the founding fathers of Keywords ethology, or animal behaviour. This simplistic description, how- Animal behaviour, ethology, stick- ever, does not provide a holistic understanding of the importance leback fish, gull, stimuli, experi- ment. RESONANCE | August 2018 845 GENERAL ARTICLE Unlike early animal of Tinbergen’s achievements, and some historical context is nec- behavioural researchers essary to appreciate this. Animal behavioural science existed who relied on a long before Tinbergen began his work on animal behaviour; the comparative psychology ff approach and viewed crucial di erence being that the predominant approach then was behaviour as a learning one of comparative psychology, and psychologists largely viewed process, Tinbergen behaviour as a learning process. Tinbergen, on the other hand, believed that the answer was more interested in understanding why “animals behave as to animal behaviour was in the hereditary they do?” and believed that the answer lay in the “hereditary influences occurring influences occurring inside the animal”. In other words, Tinber- inside the animal. gen saw behaviour as an evolutionarily adaptive trait and empha- sised a more objective, biological approach to understanding be- haviour. To understand why animals behave the way they do, Tinbergen observed animal behaviour in the natural environment of the species and conducted painstakingly designed, simple field experiments to test out his hypotheses about how animals reacted to exter- nal stimuli. He studied a variety of species ranging from wasps and stickleback fish to gulls, blackbirds and kittiwake and showed how natural observation techniques can be married successfully to experiments within laboratories. Tinbergen’s field experiments were elegantly designed simple setups that carefully teased apart functional explanations for particular behaviours exhibited by an- imals and the nature of external stimuli that may elicit certain be- havioural responses from animals. The design of his earliest ex- periments on the orientation behaviour of the beewolf, Philanthus triangulum, highlighted what was to be the hallmark of his re- search – a careful, dogged approach that incrementally addressed the effect of one variable at a time, so that it was absolutely clear what kind of cues were being used by the beewolf females to ori- ent themselves (for a better idea about the design of Tinbergen’s beewolf experiments, please see Prof. Gadagkar’s article on ani- mal behaviour experiments in this issue). Tinbergen refined this research approach throughout his career. For example, when Tinbergen started studying the reproductive behaviour of the stickleback fish in the laboratory he set up at Leiden University, he was initially interested in observing the 846 RESONANCE | August 2018 GENERAL ARTICLE courtship behaviour of male sticklebacks and carefully docu- To prove that it was the mented the many steps in their complex mating ritual. He noticed colour red that acted as that courting males turned red on their underside and attacked the ‘sign stimulus’ or ‘releaser’, Tinbergen other coloured individuals that approached their territories. The designed a set of colour red appeared to be a stimulus that set off aggressive reac- experiments wherein he tions in the males, for male sticklebacks attacked the sides of the created some rough aquarium even when a red mail van passed beside the window of models of sticklebacks, painted them in various the aquarium. To prove that it was the colour red that acted as the colours, including red, ‘sign stimulus’ or ‘releaser’, Tinbergen designed a set of exper- and presented them to iments wherein he created some rough models of sticklebacks, the male sticklebacks in painted them in various colours, including red, and presented the tank. Red coloured models always elicited them to the male sticklebacks in the tank. Red coloured models more reaction from always elicited more reaction from courting males, demonstrat- courting males, ing that it was the colour red (rather than the shape/identity of demonstrating that it was the fish) that acted as the ‘releaser’ for a specific action from the the colour red (rather than the shape/identity of male. Tinbergen then continued with his experiments to show that the fish) that acted as the particular shape, size or type of body movement can elicit certain ‘releaser’ for a specific reactions from males and females in the breeding period. For in- action from the male. stance, females will follow a red dummy fish and even try to move into a nest in the sand (where none exists), when the dummy fish is poked into the sand. Again, females will begin to lay eggs, when they are tapped repeatedly on the base of the tail (in a par- ody of the male prodding her tail base) although they observed the removal of the red fish that led them into the nest. These find- ings led Tinbergen to conclude that sticklebacks (and many other species) respond “simply to ‘sign stimuli’, i.e., to a few charac- teristics of an object rather than to the object as a whole....it is the signal, not the object, that counts. It seems to be typical of innate behaviour, and many social relationships in animals apparently are based on a system of signs.” Tinbergen’s careful experimentation (in natural conditions and in the laboratory) shows that he was an exceptional scientist, but it was his constant ability to learn that marks him as a truly out- standing biologist. Tinbergen’s thinking about animal behaviour constantly evolved during the course of his research career and he built on the findings of his earlier studies to ask newer and more RESONANCE | August 2018 847 GENERAL ARTICLE exciting questions. For example, his earlier studies such as his work on Philanthus homing behaviour, and stickleback mating behaviour were built around the central question: What stimuli elicit innate and fixed behaviour patterns in animals? In his later studies, he moved on to ask: What is the function of behaviour? His work on eggshell removal by the black-headed gull typifies this approach. In a very curious Black-headed gulls remove eggshells from the vicinity of their behaviour, black-headed nests almost immediately after hatching. Except for the top end, gulls remove eggshells shells do not markedly differ from eggs. Yet, eggs are never re- from the vicinity of their nests almost moved from the nest. Tinbergen asked the question: What is immediately after the function of the eggshell removal behaviour? He hypothesised hatching. Except for the that it was to increase the survival chances of the chicks and pro- top end, shells do not ceeded to conduct a series of experiments to test out this hypoth- markedly differ from eggs. Yet, eggs are never esis. Tinbergen observed that the insides of the eggshells were removed from the nest. white which could attract the attention of predators. To build ev- idence for this point, he and his collaborators first conducted a set of three experiments wherein they investigated if the natural colour of the black-headed gulls’ eggs acted as camouflage and thereby protected them from the attention of predators. To do this, they painted some gull eggs white and left them alongside some unpainted eggs in shallow depressions in the ground that were similar to black-headed gulls’ nests. Comparative rates of preda- tion on the experimental eggs showed that the natural colour of the gulls’ eggs made them less vulnerable to predation than those painted white. Tinbergen then went on to test if the presence of the eggshells in the vicinity of nests increased chances of preda- tion. He did this by comparing the rates of predation on gull eggs that were placed near eggshells and those that were not. He also conducted another experiment wherein eggshells were placed at increasing distances from eggs to investigate the effect of prox- imity of eggshells on egg/chick predation.
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