Orientation of locomotor behaviour is usually categorized as either kinesis or taxis. In kinesis, as previously explained, an animal’s body is not oriented in relation to a sensory stimulus; rather, the stimulus causes an alteration in speed or direction of movement. In wood lice, for example, the kinetic response alters only the rate of movement. Because wood lice tend to aggregate in moist areas, their ambulatory activity increases or decreases as the relative humidity decreases or increases, respectively. In the planarian (an aquatic, ciliated flatworm), on the other hand, the kinetic response affects only the rate at which the planarian changes its direction. Because planaria tend to stay in or return to darker areas, an increase in light intensity causes an increase in their turning responses. Generally, however, animals tend to alter both direction and speed as a single kinetic response.

In taxis, an animal orients itself in a specific spatial relationship to a stimulus. The orientation may be simply an alteration of body position or it may be an alteration of locomotor direction so that the animal moves toward, away from, or at a fixed angle to the source of the stimulus. Sources that elicit a taxis response, which may cause a modification of speed, direction, or both, seem to encompass the entire range of environmental stimuli, such as gravity (geotaxis), temperature (thermotaxis), light (phototaxis), or chemicals (chemotaxis). If the response is negative, the animal moves away from the source; if it is positive, the animal moves toward the source.

The control of the response to a taxis is of two types. In open-system control, the initial response to a stimulus has no effect on subsequent responses to the same stimulus. A male firefly, for example, locates a female by the latter’s brief flashes of light. When a male sees a female’s flash, the male turns in the direction of the female, even though the source is no longer visible. If another female flashes, however, the male responds to the second flash in exactly the same manner as it did to the first. In close-system control, on the other hand, the response is progressively altered by feedback so that all subsequent responses are adjusted to the initial response. A bat chasing a flying insect will alter its flight path to intercept that of the insect. The bat’s initial change in direction is only a general alteration of its course, but, as it approaches the insect, the bat constantly modifies its course to obtain an accurate interception.

Steering

Animals obtain accurate directional response (steering) by changing their propulsive response. Because steering relies heavily on continuous feedback (the communication cycle in which the motor output, or behaviour, is constantly being modified by the sensory input, or stimulus), it requires a precise integration of the central and peripheral nervous systems. (The central nervous system—in vertebrates, the brain and spinal cord—is that part of the nervous system that receives sensory impulses and sends out motor impulses; the peripheral nervous system consists of all the nerves that carry impulses between the central nervous system and other parts of the body.) Exteroceptive stimuli (those that originate outside the body) received by the peripheral nervous system establish the animal’s spatial position in the environment; proprioceptive stimuli (those that originate inside the body), also received by the peripheral nervous system, establish the relative position of the body units to each other. Through integration of these two sets of stimuli, the central nervous system continuously adjusts the contraction of the motor units (e.g., muscles) to obtain the desired orientation.

During locomotion, steering is a continual process. The direction of movements must be constantly adjusted to counteract environmentally produced deviations of direction. The apparently simple act of a bird flying from a tree to the ground illustrates the complexity of directional control. As the bird flies to the ground, it must be constantly aware of its height above the ground, the orientation of its body axis relative to the ground, deviations in flight direction resulting from air currents, and its speed of fall. All these parameters are determined primarily by exteroceptive stimuli received through the eyes and inner ears. The downward flight is constantly adjusted in response to these exteroceptive stimuli, and the fine control necessary for these adjustments is obtained by proprioceptive feedback.