50 Motivation
such as approach behavior. These cues also shorten copulatory
latencies (Domjan, et al., 1986). Thus, discrete Pavlovian CSs,
such as red lights, buzzers, or inanimate objects, can elicit re-
sponses that facilitate reproductive behaviors in the quail.
Contextual cues may also contribute to reproductive sig-
naling. Domjan et al. (1989) reported that male quails attempt
to mate with models of a female quail only if they have pre-
viously copulated with a live quail in the test chamber. Thus,
the location or context of previous sexual experience can act
as a signal that facilitates the occurrence of sexual behavior.
Additionally, contextual cues increase the male quail’s sperm
production (Domjan, Blesbois, & Williams, 1998). Notably,
this demonstrates that Pavlovian learning may directly en-
hance reproductive success by facilitating the bird’s ability to
fertilize multiple eggs and produce offspring.
Sexual learning also directly influences mate selection.
For example, when an orange feather is repeatedly paired
with a sexual encounter, male quails display a preference for
birds adorned with this cue. Males both spend more time near
and display more copulatory behaviors toward these females
compared to controls (Domjan, O’Vary, & Greene, 1988).
Thus, Pavlovian conditioning sways attractiveness, thereby
influencing mate selection.
Along with neutral cues, learning also facilitates the sex-
ual efficacy of sign stimuli. For example, the model of a
female’s head and neck elicits copulatory behavior in experi-
enced, but not in sexually naive, male quails (Domjan et al.,
1989). Thus, during sexual encounters these birds may learn
to identify species-typical cues, such as the plumage of
female conspecifics.
Organization of the Sexual Behavior System
Sexual behavior does not begin and end with the act of copu-
lation. Instead, species exhibit numerous behaviors that con-
tribute to reproductive success that are not directly connected
to the sex act. For example, male blue gouramis build nests
used for spawning prior to contact with female conspecifics.
This behavior improves reproductive success because nest
occupancy increases the probability that these fish will attract
a mate. Concurrently, these fish compete with male con-
specifics to secure suitable nesting areas, and they display ag-
gressive territorial behavior to defend or take control of a nest
site. Thus, because these behaviors can greatly increase re-
productive opportunities, sexual behavior can be linked to ac-
tivities that are temporally distant from the sex act.
Domjan and associates (e.g., Domjan & Hall, 1986) de-
scribed a set of behaviors that contribute to the reproductive
success of Japanese quails. Males engage in general search
behavior when they encounter cues distal to the female. For
example, birds pace around the test chamber when they en-
counter a cue that has been conditioned with a long CS-US
interval. This cue is relatively distal to the female because it
signals that a female will appear only after a long time period
elapses (Akins, Domjan, & Gutierrez, 1994). In contrast,
cues conditioned with a short CS-US interval elicit focal
search behavior. For instance, birds approach a red light that
has previously been paired with a sexual encounter (Akins
et al., 1994). This cue is relatively proximal because it signals
that the female will appear after a short time period elapses.
Male quails also engage in copulatory or consummatory sex-
ual responses (Figure 2.5). These responses are elicited by
cues signaling that a sexual encounter is imminent. Thus, fe-
male conspecifics or sign stimuli elicit copulatory behavior.
Domjan and his colleagues have characterized a range of
stimuli that elicit an array of sexual responses in the Japanese
quail. With these observations Domjan has articulated a be-
havioral systems account of sexual behavior that contains
both a stimulus and a response dimension. Each dimension
includes three categories. The response dimension includes
general search behavior, focal search behavior, and copula-
tory behavior. The stimulus dimension includes contextual
cues, local cues, and species-typical cues.
In the model, stimuli are arranged on a temporal and spa-
tial continuum that varies by the cue’s proximity to the fe-
male quail. This continuum is similar to the spatiotemporal
organization hypothesized by Timberlake (1983) in his feed-
ing behavior system and by Fanselow (1989) in his descrip-
tion of defensive behavior, both discussed earlier. Prior to
sexual conditioning, contextual and local cues are distal from
the female and do not activate sexual behavior, whereas
species-typical cues are more proximal and can elicit sexual
behavior unconditionally. After a sexual conditioning event,
contextual and local cues may elicit sexual behavior, and
responding to species-typical cues is facilitated. Thus, ac-
cording to Domjan’s view, “conditioning serves to increaseFigure 2.5 Two Japanese quail display mounting, one component of
copulatory behavior.