Comprehensive Physiology Wiley Online Library

The place of Behavior in Physiology

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Behavior as a Physiological Response
2 The Uving Being is an Open System
2.1 Behavioral Control of Inflows
2.2 Behavioral Control of Outflows
3 The Various Behaviors
3.1 Postural Adaptation
3.2 Migration
3.3 Building Microenvironments
3.4 Operant Behavior
3.5 Parental Behavior
3.6 Behavioral Self‐Adjustment
3.7 Ultimate Behavioral Self‐Adjustment: Autostimulation of the Central Nervous System
4 Efficacy and Precision of Behavior
4.1 Short‐Term Adjustments
4.2 Long‐Term Adjustments
5 The Ranking of Priorities
5.1 Complementarity of Autonomic and Behavioral Responses: The Gain of Freedom
5.2 Conflicts of Motivation: The Behavioral Final Common Path
6 Optimal Behavior
7 Conclusions
Figure 1. Figure 1.

Relative contributions of autonomic and behavioral responses in reducing threats of body temperature variation when humans are exposed to potentially hazardous conditions of heat and cold. Left: Temperatures of environments that humans could possibly encounter. Right: Thermoregulatory responses in relation to their capabilities and ranges of internal body temperature and thermal comfort. Human freedom to move about on the earth would be limited without the behavioral response

from Hardy
Figure 2. Figure 2.

Living animals are analogous to the above tank. They are open systems in steady‐state equilibrium receiving a continuous inflow and losing an equal outflow of matter and energy. Black box in upper right describes the system. Block diagram below analyzes the system with its input and output faucets related by the level h, which is a derivative of inflow and an integral of outflow

from Cabanac and Russek
Figure 3. Figure 3.

Steady state is now equipped with a sensor (float) of the regulated variable (h), a negative feedback loop on the input faucet, and a positive feed‐forward loop on the output faucet. Both loops are behavioral

from Cabanac and Russek
Figure 4. Figure 4.

Calls of American white pelican embryos at the pipped egg stage when exposed in vitro to various temperatures for 10 min. Horizontal line: mean; open box: ± 1 SE; vertical line: range; N = 15 eggs, each tested under all conditions. Embryos called for parental rescue proportionally to the deviation of temperature above and below 37.8°C

from Evans
Figure 5. Figure 5.

A pigeon standing in an experimental chamber receives a grain when it stays on the correct side of the chamber as indicated by colored lights. The rhythm of lights on and off was variable. The figure gives the logarithm of the ratio of time spent on the left of the chamber plotted as a function of the logarithm of the ratio of number of grains received on the left to number of grains received on the right during an experimental session. Symbols give the animal's behavior. The solid line is the linear regression (equation in lower right corner). The dashed line has a slope of one and passes through the origin; it represents the performance of perfect matching. It can be seen that the bird's behavior was almost perfect

from Baum and Rachlin
Figure 6. Figure 6.

Left: Some regulated variables of the organism ranked from top to bottom in order of priority, defined by the time constant tolerated before correcting a perturbation. Right: Non‐exhaustive list of somatic functions. Regulation of variables on left achieved by control (modulation) of functions on right. Heavy arrows: tight control; light arrows: loose control; dashed arrows: hypothetical control. Behavioral responses participate in defense of most regulated variables of the organism

from Cabanac and Russek
Figure 7. Figure 7.

A pigeon stands in a climatic chamber, the temperature of which (load temperature) is imposed by the experimenter. The bird can peck at a key and thus obtain a burst of cool air. The figure gives the mean resulting body temperature and behavior of three pigeons performing in 43 sessions. A: Rate of pecking: RF, respiratory frequency. The animals’ behavior was proportional to ambient temperature and therefore directly thermoregulatory. As a result, the birds saved evaporative heat loss and did not hyperventilate while maintaining stable body temperatures (B); Ta,x, axillary temperature; Ts, dorsal skin temperature; Ta temporal mean ambient temperature

from Schmidt
Figure 8. Figure 8.

Two groups of Merino wethers can feed and drink at two locations separated by the various distances indicated on the abscissa; water intake, food intake, number of shuttle trips between food and water, and resulting distance walked were monitored. A: Two drinks per day; B: Three drinks every 2 days; C: one drink per day. It can be seen that the animals modulated behavior to maintain approximately constant water intake, food intake, and cost measured by walking distance

from Squires and Wilson
Figure 9. Figure 9.

A subject walks at a steady rate of 3 km·h−1 on a treadmill in a climactic chamber. He can adjust either the slope of the treadmill or the ambient temperature. When the subject adjusts the slope, ambient temperature is imposed by the experimenter. When the subject adjusts ambient temperature, the slope is imposed by the experimenter. Each symbol shows the subject's state at the end of a 1 h session. Results obtained when slope was imposed were not different from results when ambient temperature was imposed. This means that the subject used the degree of freedom left to him to reach, with his behavioral choice, the same physiological state. In addition, results show that with one behavioral degree of freedom the subject managed to produce external work, that is, heat production, that was inversely proportional to ambient temperature and to curb increases in both heart rate and core temperature. Behavior was therefore optimal as seen from the point of view of temperature regulation

from Cabanac and Leblanc
Figure 10. Figure 10.

Diagram illustrating the phenomena and disciplines of the brain and behavioral sciences. The sciences (left column) are ranked from bottom to top in order of increasing complexity of the phenomena studied (center column). The understanding of behavior is the understanding of whole organisms

from Toates


Figure 1.

Relative contributions of autonomic and behavioral responses in reducing threats of body temperature variation when humans are exposed to potentially hazardous conditions of heat and cold. Left: Temperatures of environments that humans could possibly encounter. Right: Thermoregulatory responses in relation to their capabilities and ranges of internal body temperature and thermal comfort. Human freedom to move about on the earth would be limited without the behavioral response

from Hardy


Figure 2.

Living animals are analogous to the above tank. They are open systems in steady‐state equilibrium receiving a continuous inflow and losing an equal outflow of matter and energy. Black box in upper right describes the system. Block diagram below analyzes the system with its input and output faucets related by the level h, which is a derivative of inflow and an integral of outflow

from Cabanac and Russek


Figure 3.

Steady state is now equipped with a sensor (float) of the regulated variable (h), a negative feedback loop on the input faucet, and a positive feed‐forward loop on the output faucet. Both loops are behavioral

from Cabanac and Russek


Figure 4.

Calls of American white pelican embryos at the pipped egg stage when exposed in vitro to various temperatures for 10 min. Horizontal line: mean; open box: ± 1 SE; vertical line: range; N = 15 eggs, each tested under all conditions. Embryos called for parental rescue proportionally to the deviation of temperature above and below 37.8°C

from Evans


Figure 5.

A pigeon standing in an experimental chamber receives a grain when it stays on the correct side of the chamber as indicated by colored lights. The rhythm of lights on and off was variable. The figure gives the logarithm of the ratio of time spent on the left of the chamber plotted as a function of the logarithm of the ratio of number of grains received on the left to number of grains received on the right during an experimental session. Symbols give the animal's behavior. The solid line is the linear regression (equation in lower right corner). The dashed line has a slope of one and passes through the origin; it represents the performance of perfect matching. It can be seen that the bird's behavior was almost perfect

from Baum and Rachlin


Figure 6.

Left: Some regulated variables of the organism ranked from top to bottom in order of priority, defined by the time constant tolerated before correcting a perturbation. Right: Non‐exhaustive list of somatic functions. Regulation of variables on left achieved by control (modulation) of functions on right. Heavy arrows: tight control; light arrows: loose control; dashed arrows: hypothetical control. Behavioral responses participate in defense of most regulated variables of the organism

from Cabanac and Russek


Figure 7.

A pigeon stands in a climatic chamber, the temperature of which (load temperature) is imposed by the experimenter. The bird can peck at a key and thus obtain a burst of cool air. The figure gives the mean resulting body temperature and behavior of three pigeons performing in 43 sessions. A: Rate of pecking: RF, respiratory frequency. The animals’ behavior was proportional to ambient temperature and therefore directly thermoregulatory. As a result, the birds saved evaporative heat loss and did not hyperventilate while maintaining stable body temperatures (B); Ta,x, axillary temperature; Ts, dorsal skin temperature; Ta temporal mean ambient temperature

from Schmidt


Figure 8.

Two groups of Merino wethers can feed and drink at two locations separated by the various distances indicated on the abscissa; water intake, food intake, number of shuttle trips between food and water, and resulting distance walked were monitored. A: Two drinks per day; B: Three drinks every 2 days; C: one drink per day. It can be seen that the animals modulated behavior to maintain approximately constant water intake, food intake, and cost measured by walking distance

from Squires and Wilson


Figure 9.

A subject walks at a steady rate of 3 km·h−1 on a treadmill in a climactic chamber. He can adjust either the slope of the treadmill or the ambient temperature. When the subject adjusts the slope, ambient temperature is imposed by the experimenter. When the subject adjusts ambient temperature, the slope is imposed by the experimenter. Each symbol shows the subject's state at the end of a 1 h session. Results obtained when slope was imposed were not different from results when ambient temperature was imposed. This means that the subject used the degree of freedom left to him to reach, with his behavioral choice, the same physiological state. In addition, results show that with one behavioral degree of freedom the subject managed to produce external work, that is, heat production, that was inversely proportional to ambient temperature and to curb increases in both heart rate and core temperature. Behavior was therefore optimal as seen from the point of view of temperature regulation

from Cabanac and Leblanc


Figure 10.

Diagram illustrating the phenomena and disciplines of the brain and behavioral sciences. The sciences (left column) are ranked from bottom to top in order of increasing complexity of the phenomena studied (center column). The understanding of behavior is the understanding of whole organisms

from Toates
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Michel Cabanac. The place of Behavior in Physiology. Compr Physiol 2011, Supplement 14: Handbook of Physiology, Environmental Physiology: 1523-1536. First published in print 1996. doi: 10.1002/cphy.cp040267