The experiments of Stewart, from the Biology Faculty, inspired Kline, from the Psychology Faculty, the idea to choose rats as animal models for his research on learning processes ( Kline, 1928 Miles, 1930). Stewart was the first to perform a quantitative rodent test of motor activity. In 1898, a Clark University colleague of Kline and Small, Colin Campbell Stewart (1873–1944), published a study on the effects of alcohol, barometric pressure and type of diet on rat daily voluntary wheel running activity, as assessed by a revolving drum connected to automatic counters recording the total number of revolutions ( Stewart, 1898). On the other hand, Kline and Small introduced behavioral tests aimed at specifically evaluating cognition. In addition to purely observational studies of development, he also used some basic behavioral tests, e.g., reflex tests or taste reactivity tests, in which only qualitative responses were recorded ( Mills, 1895, 1898). Applying this method, he studied systematically the physical and psychological development of guinea pigs through a daily monitoring in a laboratory setting. Mills was the first, in the mid 1890's, to introduce for rodents the ontogenetic diary method, which consisted of observing and describing step by step the different developmental stages of a species, starting from the day of birth ( Mills, 1895). A systematic and quantitative study of rodent behavior in laboratory had yet to come. Two important elements of scientific testing were absent: systematic observations (i.e., observations at pre-set time-points according to a specific rationale) and choice of one or more quantitatively measurable behavioral outcomes as variables of interest. Although this was a first step for the study of rodent behavior in a controlled environment, the report with the findings was only anecdotal and appeared inserted within a paper on the behavior of squirrels in the wild. Mills also observed the behavior of two squirrels he captured and kept for few months, reporting, for example, how ethologically relevant behaviors could be observed, like nest-building and food storing, and how one of them learned to eat from his hand and enjoyed running on a running wheel that was installed in its home-cage ( Mills, 1888). These studies were preceded by ethological, purely observational, studies on rodent behavior in the wild, e.g., Mills's studies on squirrel behavior ( Mills, 1888, 1890, 1893), but it was only in the period 1895–1900 that behavioral tests for studying rodent behavior and psychology in a laboratory setting started to be designed. The longer is the latency, the stronger is the memory of the past electric shock exposure.īehavioral testing of rodents in a laboratory setting started in the 1890's with the studies of Thomas Wesley Mills (1847–1915) from McGill University (Montreal, Canada) and of Linus Ward Kline (1866–1961) and Willard Stanton Small (1870–1943), both from Clark University (Worcester, Massachusetts, USA) ( Mills, 1895, 1898 Kline, 1899a, b Small, 1899, 1900, 1901). In this test, the motivation is fear, the observable behavior is avoidance of a dangerous (dark) zone, and the quantifiable outcome is the latency to enter the dangerous zone, which thus serves as an index of memory. If the animal remembers receiving the shock, the dark zone should now be perceived as dangerous and hence avoided. After the initial exposure to this apparatus (the training), the animal is released a second time in the same apparatus for a memory test, but this time without the shock deliverer being active. When the animal enters the dark zone, it receives an electric shock. Being naturally photophobic and preferring dark areas, mice and rats will rapidly move from the well-illuminated zone into the dark zone, a behavior that in the wild is useful to avoid being seen by predators. Behavioral tests have three key elements: (1) a motivating factor (what motivates the animal in the test) (2) an observable behavior (which behaviors we may expect the animal exhibits in response to the test) (3) a measurable outcome (a quantifiable variable associated with the behavioral response).įor example, in the rodent step-through passive avoidance test (see d'Isa et al., 2014 for a brief history of the test), the animal is released into a strongly illuminated chamber connected to a dark zone.
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