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Recent evidence suggests a role for corticotropin-releasing factor CRF in the regulation of pair bonding in prairie voles.

Monogamous prairie and pine voles have significantly lower levels of CRF receptor type 1 CRF 1and significantly higher levels of type 2 CRF 2 binding, in NAcc than non-monogamous meadow and montane voles.

Here, we report that microinjections of CRF directly into the NAcc accelerate partner preference formation in male prairie voles. Likewise, CRF injections into NAcc of non-monogamous meadow voles also did not facilitate partner preference. Taken together, our results reveal a novel role for accumbal CRF systems in social behavior. The corticotropin-releasing factor CRF system is involved in the neurobiology underlying stress and anxiety, but much less is known about its role in social behavior.

Microtine rodents exhibit diverse social organizations and thus offer Two kinky males hotguypics ca excellent comparative approach in the study of the neurobiology of social behavior Young and Wang, Prairie Microtus ochrogaster and pine voles Microtus pinetorum are monogamous; adult mates form long-lasting selective pair bonds in the field and the laboratory Getz, Carter, and Gavish, ; Salo, Shapiro, and Dewsbury, In contrast, closely-related meadow Microtus pennsylvanicus and montane voles Microtus montanus are promiscuous and solitary Gruder-Adams and Getz, ; Shapiro and Dewsbury, Past research has revealed that the brain distribution of neuropeptide receptors for oxytocin and vasopressin appears to be responsible for the species differences in social organization Insel and Shapiro, ; Insel, Wang, and Ferris, ; Lim, Wang, Olazabal, Ren, Terwilliger, and Young, b.

However, more recent evidence Two kinky males hotguypics ca that another neuropeptide system, CRF, also appears to modulate pair bonding in prairie voles DeVries, Guptaa, Cardillo, Cho, and Carter, There are relatively few studies examining the role of stress hormones in social behavior.

One study found that administration of exogenous corticosterone to male prairie voles facilitated pair bond formation with a novel female DeVries, DeVries, Taymans, and Carter, A subsequent study found that CRF administered intracerebroventricularly i. Furthermore, partner preference was blocked by i. These data suggest that CRF may play a role in pair bond formation via anxiety-independent mechanisms through the involvement of centrally acting brain receptors. Since the CRF system has been implicated in the regulation of pair bond formation, we predicted that neural circuits for this system would differ between monogamous and non-monogamous species.

Receptor binding appeared to correlate with monogamous social structure in several brain regions; however, only the nucleus accumbens NAcc consistently segregated with both monogamous vole species and both non-monogamous vole species.

First, we determined whether CRF injected directly into NAcc can facilitate partner preference formation after an abbreviated cohabitation time with the partner. Next, we performed the identical experiment in non-monogamous meadow voles. The results from these studies demonstrate for the first time that CRF, acting in the NAcc can promote social attachment, and furthermore, that both CRF 1 and CRF 2 receptors are involved in this process.

After weaning at 21 days of age, subjects were housed in same-sex sibling pairs or trios and water and Purina rabbit chow provided ad libitum. All cages were maintained on a Two kinky males hotguypics ca from 87 male prairie voles were included in the CRF pharmacology experiments, along with equal numbers of stimulus female prairie voles for the pair bonding assay.

Data from 10 male meadow voles were used as well, along with equal numbers of stimulus female meadow voles.

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Subjects were anesthetized with sodium pentobarbital 2. After 3—5 Two kinky males hotguypics ca recovery, subjects received microinjections nl per side of either artificial CSF or drug dissolved in CSF. Injections were made with a 33 gauge needle that extended 1 mm below the guide cannula into the target area. Animals were divided into one of four groups: Each animal received bilateral injections of nl volume prior to an abbreviated 6 hour cohabitation with a novel female.

The concentration of 0. Cohabitation for 6 hours without mating consistently does not induce partner preference, as demonstrated in previous studies Aragona et al. Immediately following cohabitation, subjects were tested for partner preference. Subjects were allowed to freely roam throughout the apparatus, and time spent in contact with the partner and stranger was quantified over the course of the 3 hour test.

Locomotor activity was measured during the partner preference test to Two kinky males hotguypics ca whether chosen doses of CRF treatment affected general locomotor activity or anxiety-like behavior, as previously described Hotta, Shibasaki, Arai, and Demura, The number of cage crossings through both tunnels of the partner preference apparatus was assessed by using infrared detectors.

There are four infrared beams across the three cages of the partner preference apparatus, with two beams flanking each tunnel connecting two cages.

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The total number of photo beam breaks was totaled for each animal during the three-hour period. After behavioral testing, subjects were sacrificed and injection sites were verified histologically.

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Animals were cannulated bilaterally into NAcc as described above and randomly Two kinky males hotguypics ca to one of two groups: Cohabitation and partner preference tests were performed exactly as described above for prairie voles. Adult male meadow voles from our colony typically do not form partner preferences when cohabitated with a female Lim et al. Adult prairie voles were cannulated bilaterally into NAcc as described above and divided into one of three groups: Anti-sauvagine was obtained from Sigma St.

Emory University, Atlanta, GA. Each animal received bilateral injections of nl volume directly into NAcc prior to an abbreviated 6 hour cohabitation with a novel female. Immediately following cohabitation, subjects were tested for partner preference as described above. Animals whose cannulation sites were placed outside the NAcc were excluded from data analysis and not reflected in the total number of animals used.

Data from the partner preference test for each experiment were analyzed using a 2 way ANOVA in which stimulus partner or stranger and treatment were factors.

Bonferroni corrections for the level of significance were made for each experiment in Two kinky males hotguypics ca to minimize risk of type-I error risk due to multiple comparisons. Males were categorized as having developed a partner preference if they spent more than twice as much time in contact with the partner than the stranger.

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Data collected for locomotor activity was totaled as the number of infrared beam breaks for each animal, and averaged within each treatment group. Adult prairie voles were deeply anesthetized between the hours of Briefly, endogenous peroxidase activity was quenched by minute incubation with 0.

For the CRF-specific antibody, blocking was performed by a five hour incubation with 4. The primary rabbit antibodies recognizing Urocortin-1 Sigma-Aldrich, St. Louis, MO were used in dilution 1: Biotynylated anti-rabbit secondary antibodies Vector Laboratory Inc.

The specificity of immunostaining was assessed by a complete lack of immunoreactivity in areas not known to express CRF or Urocortin Futhermore, control preabsorption experiments for these antibodies were previously performed Bachtell, Weitemier, Galvan-Rosas, Tsivkovskaia, Risinger, Phillips, Grahame, and Ryabinin, Two kinky males hotguypics ca image analysis was performed using a system consisting of an Olympus BX40 microscope and a high-resolution digital videocamera Olympus Qcolor3 interfaced to a Macintosh personal computer running OS-X.

Images from a single section best matched across animals for each brain region were digitally collected at the same lighting intensity. Since no immunopositive staining Two kinky males hotguypics ca observed in cell bodies in NAcc, the number of immunoreactive neurons was not quantitatively counted.

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It had been previously demonstrated that infusion of CRF i. The earlier study showed that dose-dependent i.

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The doses that we used, 0. Prairie voles injected with the lower CRF dose, 0. In contrast, prairie voles receiving bilateral injections of the fold higher CRF dose, 0. Furthermore, while only Two kinky males hotguypics ca of the 13 control animals displayed a partner preference, defined as spending twice as much time in contact with the partner compared to the stranger, 12 of the 15 animals receiving 0.

The 3 animals that did not display a partner preference had strong stranger preferences, which likely contributed to the lack of a main effect of treatment or an interaction effect in the 2-way ANOVA. Thus, while CRF infusions into the NAcc significantly increased the time in contact with the partner relative to the stranger, it did not result in an overall increase in contact time with the partner.

Animals that received 0. Animals receiving a fold higher dose, 0.

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Partner preference is defined as spending twice as much time in contact with the partner than with the stranger. Indeed, meadow voles injected with CSF, or the high dose 0. Bilateral micro-injections of CRF into NAcc fail to facilitate partner preference in non-monogamous male meadow voles.

Adult meadow voles injected with artificial CSF, or 0. Prairie voles injected with a cocktail of 0.

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Interestingly, prairie voles injected with a cocktail of 0. Control prairie voles injected with 0.

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In addition, while 8 out of the 10 CRF-treated prairie voles displayed a partner preference, only 11 of the 25 animals receiving CRF 1 antagonist, and 6 out of 15 animals receiving CRF 2 antagonist, displayed a partner preference Figure 3B. A Animals injected with 0. Results are shown in Table 1.