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Oxytocin

Social Behavior Network

Chapter 1: Oxytocin

Chapter 2: Social Behaviour Network

amygdala and the medial bed nucleus of stria terminalis,BSTm), the lateral septum (LS), the preoptic area (POA),the anterior hypothalamus (AH), the ventromedial hypo-thalamus (VMH), and the midbrain (Fig. 1 and Table 1).The relevant midbrain areas include the periaqueductal gray(PAG) and various areas of the tegmentum that linkforebrain regions with motoneuron pools of the hindbrain.The other five nodes lie within the basal (‘‘limbic’’)forebrain. Newman (1999) proposed that these areascomprise a social behavior network based upon a fewimportant criteria. First, each of the nodes has beenimplicated in the control of multiple forms of socialbehavior. These include aggression, appetitive and con-summatory sexual behavior, various forms of communica-tion, social recognition, affiliation, bonding, parentalbehavior, and responses to social stressors (Bamshad andAlbers, 1996; Coolen et al., 1997; Cushing et al., 2003;Delville et al., 2000; Ferguson et al., 2002; Gammie andNelson, 2001; Heeb and Yahr, 2001; Kalinichev et al., 2000;Kirkpatrick et al., 1994; Kollack-Walker and Newman,1995; Kollack-Walker et al., 1997; Lim and Young, 2004;Lonstein et al., 1998; Morgan et al., 1999; Sheehan et al.,2001; Wang et al., 1997). The nodes are also bidirectionallyconnected (Coolen and Wood, 1998; Coolen et al., 1998;Dong and Swanson, 2004; Risold and Swanson, 1997b),and each area contains sex steroid receptors that are

essential for the sexual differentiation and temporalcoordination of social behavior (Commins and Yahr,1985; Morrell and Pfaff, 1978; Simerly et al., 1990; Woodand Newman, 1995). The mammalian brain obviouslycontains a large number of other areas that are relevant forsocial behavior (e.g., other basal forebrain areas thatregulate stress and reward processes, and cortical areas thatserve executive functions); thus Newman’s network shouldbe regarded as the ‘‘core’’ of the social brain, not the socialbrain in toto.

Newman (1999) proposes that this network of brain areasdoes not contain segregated, linear systems for the regulationof each kind of social behavior. Rather, each node of thenetwork responds to a variety of stimuli, with each socialcontext and behavioral response being associated with adistinct pattern of response across the nodes. For instance,some of the same areas show increases in immediate earlygene (IEG) activity during male sexual behavior, femalesexual behavior, and male aggression, but the overall patternis distinct for each behavioral context (Fig. 1). Although thismodel is in some ways very simplified (e.g., each area mayhave distinct neuronal populations with different responseprofiles), the idea is nonetheless compelling and supportedby a good body of data (see references above).

Increasing evidence suggests that this network is presentin all vertebrates. While most relevant findings have comefrom birds, consistent data are also available for amphibians,fish, and reptiles (see Table 1 and the following twosections), and Newman’s model has been explicitly appliedas a conceptual framework for data in geckos (Eublepharismacularius) (Crews, 2003). Interestingly, data from geckosdemonstrate that behavioral variations are correlated withdistinct patterns of functional connectivity within thenetwork (Sakata and Crews, 2004; Sakata et al., 2000), afinding that lends good empirical support to Newman’s(1999) ideas about the significance of distributed activationpatterns.

Our own research expands upon these studies in twoimportant respects: First, our work on the vocal circuitry ofthe plainfin midshipman fish (Porichthys notatus) providesthe first comprehensive mapping of the network’s con-nections in a non-mammal and has yielded strong evidencethat the social behavior network arose in the earliestvertebrates (Goodson and Bass, 2002). As detailed below,the vocal system of the midshipman offers opportunitiesfor systems-level experiments on the network that are notpossible in most other animals (i.e., the whole brain can beexposed during analyses of social behavior patterning),thus understanding the comparative organization of therelevant circuitry in fish is particularly valuable. Secondly,our anatomical and behavioral experiments in both fishand birds demonstrate that many functional, structural, andneuroendocrine features are exceptionally similar to thosein mammals, thereby rendering detailed comparisonsacross classes much more feasible than we initiallyanticipated.

Fig. 1. The social behavior network as originally suggested for mammals(schematics modified from Newman, 1999). The network is comprised ofsix nodes—the extended medial amygdala (i.e., the medial amygdala andthe medial bed nucleus of stria terminalis), the lateral septum (LS), thepreoptic area (POA), the anterior hypothalamus (AH), the ventromedialhypothalamus (VMH), and various areas of the midbrain, including theperiaqueductal gray. Each of the nodes binds sex steroid hormones and hasbeen implicated in the control of multiple forms of social behavior.Newman (1999) proposes that this network does not contain segregated,linear systems for each kind of behavior. Rather, as shown in theseschematic representations of immediate early gene data, each behavioralcontext is associated with a distinct pattern of activation across the nodes.

J.L. Goodson / Hormones and Behavior 48 (2005) 11–2212

Violet-eared waxbill

Melba finchMelba finch

Spice finch

Angolan blue waxbill

Zebra Finch

Chapter 3: The Zebra Finch

Breeding flocks

Foragingflocks

Ripening of grass seeds

Chapter 4:

The target article

References and Notes1. F. d’Errico et al., J. Hum. Evol. 4

8, 3 (2005).

2. C. S. Henshilwood et al., Science 295, 1

278 (2002).

3. C. W. Marean et al., Nature 449, 90

5 (2007).

4. S. McBrearty, A. S. Brooks, J. Hum.

Evol. 39, 453 (2000).

5. C. A. Tryon,S. McBrearty, J. Hum.

Evol. 42, 211

(2002).

6. M. Lombard, J. Hum. Evol. 53, 40

6 (2007).

7. J. D. Clark,J. World Prehist. 2, 235 (1988).

8. C. S. Henshilwood et al., J. Hum. Evol. 4

1, 631 (2001).

9. S. H. Ambrose, K. G. Lorenz

, in The Emergence of Modern

Humans: An Archaeological Perspective, P. Mellars, Ed.

(Edinburgh Univ. Press, Edinburgh, 1990)

, pp. 3–33.

10. H. V. Merrick, F. H. Brown, W. P

. Nash, in Society, Culture,

and Technology in Africa, S. T. Childs, Ed.(Univ. of

Pennsylvania Museum of Archaeology

and Anthropology,

Philadelphia, PA, 1994), pp. 2

9–44.

11. Z. Jacobs et al., Science 322, 733 (2008).

12. C. Tribolo et al., J. Archaeol. Sci. 36, 730 (2009).

13. T. P. Volman, in Southern African Prehistory and

Paleoenvironment, R.G. Klein, Ed. (B

alkema, Rotterdam,

Netherlands, 1984), pp. 169–2

20.

14. A. J. Mackay, Archaeol. Sci. 35, 614 (2008).

15. H. J. Deacon, in The Human Revoluti

on, P. Mellars, C. Stringer,

Eds. (Princeton Univ. Press, Prince

ton, NJ, 1989), pp. 547–564.

16. S. Wurz, S. Afr. Archaeol. Bull. 54, 38

(1999).

17. G. S. McCall, J. Archaeol. Sci. 34

, 1738 (2007).

18. M. Hanckel, Arch. Oceania 20, 98 (1985).

19. J. A. Webb, M. Domanski,

Archaeometry 50, 555(2008).

20. Materials and methods are available as supporting

material on Science Online.

21. A. W. Pelcin, J. Archaeol. Sci. 24

, 749 (1997).

22. A. Goudie,Prog. Phys. Geogr. 30

, 703 (2006).

23. M. Fener et al., Rock Mech. Rock Eng. 38,

329 (2005).

24. E. Callahan, Arch. Eastern North America 7, 1 (1979).

25. J. J. Shea,J. Archaeol. Sci. 33, 8

23 (2006).

26. A. I. R. Herries, in Australian Archaeometry, A. Fairb

rain,

S. O'Conner, Eds. (Australian National Univ.

Press,

Canberra, Australia, 2009), pp

. 235–253.

27. M. J. Aitken, Thermoluminescence

Dating (Academic Press,

London, 1985).

28. J. J. Flenniken, J. P. White,

Aust. Aborig. Stud. 1, 43 (1983).

29. J. D. Clark,J. W. K. Harris, Afr. Arch

aeol. Rev. 3, 3 (1985).

30. N. Goren-Inbar et al., Science 30

4, 725 (2004).

31. We thank the Institute for S

ocial Science Research staff at

ASU and the MosselBay Archaeology

Project staff fortheir

assistance, the Dias Museum for field facilitie

s, South African

Heritage Resources Agency and

Heritage Western Cape for

permits, and Iziko Museum for providing the

Blombos Sands

Still Bay point. This research was

funded by NSF (USA)

(grants BCS-9912465, BCS-0130713, an

d BCS-0524087to

C.W.M.); the Hyde Family Trust; a

nd the Institutefor Human

Origins, ASU. Additional support

to A.I.R.H. was provided

by M. Hill at theUniversity of Liv

erpool Geomagnetism

Laboratory, andfunds were prov

ided by the Faculty of

Medicine at UNSW and Australian R

esearch Councilgrant

DP0877603. M.C.M. was funded

via a Marie Curie

International Outgoing fellowshi

p (PIOF-GA-2008-21994).

Supporting Online Material

www.sciencemag.org/cgi/conten

t/full/325/5942/859/DC1

Materials and Methods

SOM Text

Figs. S1 to S7

Tables S1 to S8

References

15 April 2009; accepted 10 July 2009

10.1126/science.1175028

Mesotocin and Nonapeptide Receptors

Promote Estrildid Flocking Behavior

James L. Goodson,* Sara E. Schrock, James D.

Klatt, David Kabelik, Marcy A. Kingsbury

Proximate neural mechanisms th

at influence preferences for gro

ups of a given size are almost w

holly

unknown. In thehighly gregariou

s zebra finch (Estrildidae: Taenio

pygia guttata), blockade of nona

peptide

receptors by anoxytocin (OT) an

tagonist significantly reduced ti

me spent with large groups and

familiar

social partners independent of t

ime spent in social contact. Opp

osing effects were produced by

central

infusions of mesotocin (MT, avia

n homolog of OT). Most drug eff

ects appeared tobe female-specif

ic. Across

five estrildid finch species, specie

s-typical group size correlates wi

th nonapeptidereceptor distribu

tions in

the lateral septum, and sociality

in female zebrafinches was redu

ced by OT antagonist infusions i

nto the

septum but not a control area. We propo

se that titrationof sociality by M

T represents a phylogenetically

deep frameworkfor the evolution

of OT’s female-specific roles in p

air bonding andmaternal functio

ns.

Sociality, as defined by modal species-

typical groupsize, is a core component

of social organization that strongly affects

reproductive behavior, disease transmission, r

e-

source exploitation, and defen

se (1, 2). However,

the neural mechanisms that titrat

e sociality and

regulate the preference to live as singletons,in

large groups, or somewhere in

between are largely

unknown. Thislikely reflects limited

tractability,

partly because the space requirements of large

species-typical group sizes may be difficult to

accommodatein experimenta

l settings and,more

importantly, because the beha

vioral dimension of

sociality is difficult to isolate in comparative

studies. For instance, becaus

e rodent species that

differ in sociality also tend to differ in mating

system, patterns of parental c

are, and other as-

pects of behavior and ecolog

y that can influence

neural and endocrine mechanisms associated

with sociality (3, 4), comparative studies a

re not

ExperimentalArchaeological

A

0246

81012

1 1.5 2 2.5 3 3.5 4 4.5 5 5.5 6 6.5

Mor

e

Unheated

Heated

0

5

10

15

20

Freq

uenc

yFr

eque

ncy

B

0

2

4

6

8

DBCS (HP)

LCMSA

Experimental Gloss

Archaeological GlossC(50/311)

(22/24)

1 1.5 2 2.5 3 3.5 4.5 5 5.5 6 6.5

Mor

e

4

GU

Fig. 3. Analysis of gloss from

experimental and archaeologica

l silcrete sam-

ples. (A) Photo of typical heat-tre

atment gloss from an experimenta

lly treated

silcrete flake (left) and an archaeologicalspecimen from PP5-6 (right). The

rougher surfacerepresents the p

re-treatment texture of the stone

surface and

the smooth rippled surface repre

sents the post–heat-treatment fr

acture plane.

(B) Histogram showing MG values in GU for unheated (black) and heated

(red)

experimental samples. The verti

cal dashed line indicates the poi

nt above which

there are no unheated samples. (

C) The experimental results are t

hen compared

to archaeological samples. MG values are provi

ded for archaeological samples

from the PP5-6 DBCS(~60 to 65 ka) a

ggregate (top) and PP13B LC-MS

A Lower

(~164 ka) aggregate (bottom).

Department ofBiology, Indian

a University, Bloomington,

IN 47405, USA.

*To whom correspondenceshould be addressed. E-m

ail:

[email protected]

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-Oxytocin -Social Behaviour Network

-Zebra finches

Familiar

informative with respect to group size. Even insongbirds, which are speciose and socially di-verse, only few opportunities exist for controlledcomparisons, most notably in the family Estrildi-dae (finches, waxbills, and munias; about 132species total). All estrildid species are monoga-mous, exhibit long-term or life-long pair bonds,and show biparental care. However, even withinspecific ecological niches, estrildids display anextraordinary diversity in sociality, rangingfrom territorial male-female pairs to groupscontaining dozens or hundreds of coloniallybreeding pairs (5). This family includes the zebrafinch (Taeniopygia guttata), a socially tractablelaboratory songbird (6).

In mammals, the neuropeptides argininevasopressin (VP) and oxytocin (OT) influencenumerous behavioral processes, includingparental care, anxiety, and monogamous pairbonding (7–11). Behavior may also be influencedin a sex-specific manner [e.g., pair bonding (12)]

such that VP is required for a given behavior inmales, but in females the same behavior is de-pendent on OT. It has been postulated that sep-arate nonapeptide clades giving rise to VP andOT originated because of a duplication of thearginine vasotocin (VT) gene in fish (13) and thatsex-specific and affiliation-related nonapeptidefunctions date back to this duplication (14–16).

The major OT and VP cell groups in themammalian brain are found in the preoptic areaand hypothalamus, and these populations arestrongly conserved across the vertebrate classes,as are their central and hypophyseal projections(3, 17). In amniotes, OT-like peptide neurons arelargely restricted to the hypothalamus, whereashomologous cell groups lie within the preopticarea of anamniotes (3). VT and VP cell groupsare found in these same areas, although tetrapodsalso produce VTand VP in numerous extrahypo-thalamic parts of the brain, including the medialextended amygdala (bed nucleus of the stria

terminalis, BSTm) (3). Most vertebrates (includ-ing birds) have multiple VTand VP receptor sub-types but only a single OT-like receptor (18).

Given that OT promotes bonding in mam-mals (7), we conducted experiments to test thehypothesis that endogenous MT promotes pref-erences for familiar conspecifics in birds (19).Zebra finches were housed in same-sex groups ofsix birds and individually tested for novel-familiar preferences by recording the amount oftime subjects spent in close proximity to fivefamiliar cagemates versus five novel same-sexconspecifics (see diagram in Fig. 1A). We estab-lished behavioral effects by using peripheralinjections of the selective OT receptor antagonistdesGly–NH2,d(CH2)5[Tyr(Me)2, Thr4] ornithinevasotocin [OTA; 5 mg subcutaneously (20)] orsaline, and, on the basis of the positive results ofthis experiment, we fitted finches with chronicguide cannulae and replicated the experiment byusing intracerebroventricular infusions of OTA[250 ng; OTA and peptide dosages were used onthe basis of previous experiments (15, 21)]. Alltests were conducted by using a within-subjectsdesign, and tests were spaced 2 days apart.

As predicted, peripheral administration ofOTA significantly reduced the percent of timethat subjects spent in close proximity to familiarcagemates (Fig. 2A). This effect is most pro-nounced in females, although significance isachieved only for the main effect of treatment[F1,1,21 = 3.833, P < 0.05, repeated-measuresanalysis of variance (ANOVA)]. Central OTAinfusions exerted a similar and significantlysex-specific effect (Fig. 2B; sex by treatmentF1,1,20 = 4.019, P < 0.05). In contrast, centraladministrations of 50 ng MT but not VT tendedto increase the time spent with familiar cagemates(Fig. 2C; main effect of treatment F1,2,20 =3.900, P < 0.05), especially for females, althoughthe main effect of treatment in this experimentappears to come from the difference between VTand MT. Despite weak trends, no treatment ef-fects were observed in any of these experimentsfor the percent of time spent with novel con-specifics (fig. S1).

We tested the hypothesis that endogenous MTpromotes sociality by using the same apparatus asabove, but, rather than being exposed to novel andfamiliar conspecifics, subjects were exposed to agroup of two same-sex conspecifics at one end ofthe testing cage and a group of 10 same-sexconspecifics at the other end (as in Fig. 1B). In thefirst two experiments, subjects received OTA orsaline subcutaneously andwere exposed to groupsof either novel conspecifics (Fig. 3A) or familiarconspecifics (Fig. 3B). For this latter experiment,stimulus animals were housed in a cage immedi-ately adjacent to the subject’s cage for a minimumof 5 days before testing. Significant sex by treat-ment interactions were obtained in both experi-ments, with OTA reducing the percent of time thatfemale subjects spent in close proximity to the largegroup (F1, 1, 21 = 8.426, P < 0.01 and F1,1,22 =4.871, P < 0.05, respectively, repeated-measures

Fig. 1. Choice apparatus design. A 1-m-wide testing cage was subdivided into zones by sevenperches (indicated by thin lines). Subjects were considered to be within close proximity when theywere within 6 cm of a stimulus cage (corresponding to the perches closest to the sides). For novel-familiar choice tests (A), the two stimulus cages contained either five familiar cagemates or fiveunfamiliar individuals (all of the same sex). For sociality tests (B), the stimulus cages containedeither 2 or 10 same-sex conspecifics. Sides were counterbalanced across subjects. Tests were 5 min,and subject location was recorded at 15-s intervals.

Fig. 2. Zebra finches were tested in a novel-familiar choice paradigm (as in Fig. 1A) after administrationsof OTA and nonapeptides. Females and males are shown in green and purple, respectively. (A) OTAdelivered subcutaneously significantly reduced time spent in close proximity to familiar same-sexcagemates (*P < 0.05, main effect of treatment; n = 12 males, 11 females). Significant pairwisecomparisons within a sex are indicated by different letters above the bars. Data plotted are mean T SE. (B)A similar result is observed after intracerebroventricular infusions of OTA, although a significant sex bytreatment interaction is obtained (single pound symbol indicates P < 0.05; n = 11 males, 11 females). (C)Central infusions of MT and VT differentially influence time spent with familiar cagemates (*P < 0.05,main effect of treatment; n = 11 males, 11 females). Corresponding data for time spent in close proximityto novel conspecifics is shown in fig. S1.

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Unfamiliar

Large(familiar)

Small(familiar)

Focal bird has been injected with saline, mesotocin, or an oxytocin antagonist.









Fig. 1. Choice apparatus design. A 1-m-wide testing cage was subdivided into zones by seven

perches (indicated by thin lines). Subjects were considered to be within close proximity when they

were within 6 cm of a stimulus cage (corresponding to the perches closest to the sides). For novel-

familiar choice tests (A), the two stimulus cages contained either five familiar cagemates or five

unfamiliar individuals (all of the same sex). For sociality tests (B), the stimulus cages contained

either 2 or 10 same-sex conspecifics. Sides were counterbalanced across subjects. Tests were 5 min,

and subject location was recorded at 15-s intervals.

Fig. 2. Zebra finches were tested in a novel-familiar choice paradigm (as in Fig. 1A) after administrations

of OTA and nonapeptides. Females and males are shown in green and purple, respectively. (A) OTA

delivered subcutaneously significantly reduced time spent in close proximity to familiar same-sex

cagemates (*P < 0.05, main effect of treatment; n = 12 males, 11 females). Significant pairwise

comparisons within a sex are indicated by different letters above the bars. Data plotted are mean T SE. (B)

A similar result is observed after intracerebroventricular infusions of OTA, although a significant sex by

treatment interaction is obtained (single pound symbol indicates P < 0.05; n = 11 males, 11 females). (C)

Central infusions of MT and VT differentially influence time spent with familiar cagemates (*P < 0.05,

main effect of treatment; n = 11 males, 11 females). Corresponding data for time spent in close proximity

to novel conspecifics is shown in fig. S1.


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Conclusion? Mesotocin increases female preference for familiar conspecitics.

Peripheral Central Central

Females Males

Fig. 3. Zebra finches were testedin a group-size choice paradigm(as in Fig. 1B) after administra-tions of OTA and nonapeptides.The percent of time spent inclose proximity to the largegroup is shown (A to D). Corre-sponding data for time spent inclose proximity to the small groupare shown directly below (E to H).Data plotted are mean T SE.Females and males are shownin green and purple, respective-ly. Significant pairwise compar-isons within a sex are indicatedby different letters above thebars. (A) OTA delivered sub-cutaneously significantly reducesthe percent of time spent inclose proximity to the large groupin sociality tests with unfamiliar social partners but only in females. (E) This

pattern is reversed for time spent in close proximity to the small group (single

pound symbol, P < 0.05; double pound symbol, P < 0.01; sex by treatment

interaction; n = 12 males, 11 females). (B and F) Similar results are obtained

with familiar social partners after subcutaneous delivery of OTA (single pound

symbol, P < 0.05, sex by treatment interaction; *P < 0.05, main effect of

treatment; n = 12 males, 12 females). (C and G) Central OTA infusions

decrease time in close proximity to the large group and increase time in close

proximity to the small group (tests conducted with unfamiliar social partners;

*P < 0.05, ***P < 0.001, main effect of treatment; n = 12 males, 12 females).

(D) Central infusions of MT but not VT produce a female-specific increase in

the percent of time in close proximity to the large group (tests conducted with

unfamiliar social partners; single pound symbol, P < 0.05, sex by treatment

interaction; n = 17 males, 16 females).

Fig. 4. [I125]-OVTA binding sites in the LS differentiateterritorial and flocking finch species. Hp indicates hippo-campus; LSc.d, dorsal zone of the caudal lateral septum;LSc.v,vl, ventral and ventrolateral zones of the caudallateral septum; N, nidopallium; PLH, posterolateral hypo-thalamus; and TeO, optic tectum. (A to C) Representativeautoradiograms of binding sites in two sympatric con-geners, the territorial violet-eared waxbill (A) and thegregarious Angolan blue waxbill (B) plus the highly gre-garious zebra finch (C). (D) Densities of [I

125]-OVTA bindingsites in the dorsal (pallial) LSc of two territorial species(melba finch, MF, and violet-eared waxbill, VEW), a mod-erately gregarious species (Angolan blue waxbill, ABW),and two highly gregarious species (spice finch, SF, andzebra finch, ZF). Territorial and flocking species are shownin blue and orange, respectively. No sex differences areobserved and sexes were pooled. Total n = 23 (SOM text).Different letters above the boxes denote significant speciesdifferences (Mann-Whitney P < 0.05) following significantKruskal-Wallis. Box plots show the median (line), 75th and25th percentiles (box), and 95% confidence interval (errorbars). (E) Binding densities tend to reverse in the subpallialLSc (Kruskal-Wallis P = 0.06), suggesting that species dif-ferences in sociality are most closely associated with therelative densities of binding sites along a dorso-ventralgradient, as confirmed in (F).

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Conclusion? Mesotocin makes female zebra finches prefer larger groups.

Peripheral, unfamiliar

Peripheral,familiar

Central, unfamiliar

Central, unfamiliar

Females Males

Violet-eared waxbill

Melba finchMelba finch

Spice finch

Angolan blue waxbill

Zebra Finch

Conclusion? Oxytocin receptor distribution varies across species. This leads to differences in “sociality”.

Fig. 3. Zebra finches were testedin a group-size choice paradigm(as in Fig. 1B) after administra-tions of OTA and nonapeptides.The percent of time spent inclose proximity to the largegroup is shown (A to D). Corre-sponding data for time spent inclose proximity to the small groupare shown directly below (E to H).Data plotted are mean T SE.Females and males are shownin green and purple, respective-ly. Significant pairwise compar-isons within a sex are indicatedby different letters above thebars. (A) OTA delivered sub-cutaneously significantly reducesthe percent of time spent inclose proximity to the large groupin sociality tests with unfamiliar social partners but only in females. (E) Thispattern is reversed for time spent in close proximity to the small group (singlepound symbol, P < 0.05; double pound symbol, P < 0.01; sex by treatmentinteraction; n = 12 males, 11 females). (B and F) Similar results are obtainedwith familiar social partners after subcutaneous delivery of OTA (single poundsymbol, P < 0.05, sex by treatment interaction; *P < 0.05, main effect oftreatment; n = 12 males, 12 females). (C and G) Central OTA infusions

decrease time in close proximity to the large group and increase time in closeproximity to the small group (tests conducted with unfamiliar social partners;*P < 0.05, ***P < 0.001, main effect of treatment; n = 12 males, 12 females).(D) Central infusions of MT but not VT produce a female-specific increase inthe percent of time in close proximity to the large group (tests conducted withunfamiliar social partners; single pound symbol, P < 0.05, sex by treatmentinteraction; n = 17 males, 16 females).

Fig. 4. [I125]-OVTA binding sites in the LS differentiateterritorial and flocking finch species. Hp indicates hippo-campus; LSc.d, dorsal zone of the caudal lateral septum;LSc.v,vl, ventral and ventrolateral zones of the caudallateral septum; N, nidopallium; PLH, posterolateral hypo-thalamus; and TeO, optic tectum. (A to C) Representativeautoradiograms of binding sites in two sympatric con-geners, the territorial violet-eared waxbill (A) and thegregarious Angolan blue waxbill (B) plus the highly gre-garious zebra finch (C). (D) Densities of [I125]-OVTA bindingsites in the dorsal (pallial) LSc of two territorial species(melba finch, MF, and violet-eared waxbill, VEW), a mod-erately gregarious species (Angolan blue waxbill, ABW),and two highly gregarious species (spice finch, SF, andzebra finch, ZF). Territorial and flocking species are shownin blue and orange, respectively. No sex differences areobserved and sexes were pooled. Total n = 23 (SOM text).Different letters above the boxes denote significant speciesdifferences (Mann-Whitney P < 0.05) following significantKruskal-Wallis. Box plots show the median (line), 75th and25th percentiles (box), and 95% confidence interval (errorbars). (E) Binding densities tend to reverse in the subpallialLSc (Kruskal-Wallis P = 0.06), suggesting that species dif-ferences in sociality are most closely associated with therelative densities of binding sites along a dorso-ventralgradient, as confirmed in (F).


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Their claim: mesotocin promotes preference for familiar conspecifics and

leads to larger group sizes.My argument: they may not be measuring what they think they’re

measuring, and there are better ways

to measure sociality.

Familiar









Fig. 1. Choice apparatus design. A 1-m-wide testing cage was subdivided into zones by sevenperches (indicated by thin lines). Subjects were considered to be within close proximity when theywere within 6 cm of a stimulus cage (corresponding to the perches closest to the sides). For novel-familiar choice tests (A), the two stimulus cages contained either five familiar cagemates or fiveunfamiliar individuals (all of the same sex). For sociality tests (B), the stimulus cages containedeither 2 or 10 same-sex conspecifics. Sides were counterbalanced across subjects. Tests were 5 min,and subject location was recorded at 15-s intervals.

Fig. 2. Zebra finches were tested in a novel-familiar choice paradigm (as in Fig. 1A) after administrationsof OTA and nonapeptides. Females and males are shown in green and purple, respectively. (A) OTAdelivered subcutaneously significantly reduced time spent in close proximity to familiar same-sexcagemates (*P < 0.05, main effect of treatment; n = 12 males, 11 females). Significant pairwisecomparisons within a sex are indicated by different letters above the bars. Data plotted are mean T SE. (B)A similar result is observed after intracerebroventricular infusions of OTA, although a significant sex bytreatment interaction is obtained (single pound symbol indicates P < 0.05; n = 11 males, 11 females). (C)Central infusions of MT and VT differentially influence time spent with familiar cagemates (*P < 0.05,main effect of treatment; n = 11 males, 11 females). Corresponding data for time spent in close proximityto novel conspecifics is shown in fig. S1.


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Unfamiliar

Large(familiar)

Small(familiar)

Focal bird has been injected with saline, mesotocin, or an oxytocin antagonist.















Fig. 4. [I125]-OVTA binding sites in the LS differentiate

territorial and flocking finch species. Hp indicates hippo-

campus; LSc.d, dorsal zone of the caudal lateral septum;

LSc.v,vl, ventral and ventrolateral zones of the caudal

lateral septum; N, nidopallium; PLH, posterolateral hypo-

thalamus; and TeO, optic tectum. (A to C) Representative

autoradiograms of binding sites in two sympatric con-

geners, the territorial violet-eared waxbill (A) and the

gregarious Angolan blue waxbill (B) plus the highly gre-

garious zebra finch (C). (D) Densities of [I125]-OVTA binding

sites in the dorsal (pallial) LSc of two territorial species

(melba finch, MF, and violet-eared waxbill, VEW), a mod-

erately gregarious species (Angolan blue waxbill, ABW),

and two highly gregarious species (spice finch, SF, and

zebra finch, ZF). Territorial and flocking species are shown

in blue and orange, respectively. No sex differences are

observed and sexes were pooled. Total n = 23 (SOM text).

Different letters above the boxes denote significant species

differences (Mann-Whitney P < 0.05) following significant

Kruskal-Wallis. Box plots show the median (line), 75th and

25th percentiles (box), and 95% confidence interval (error

bars). (E) Binding densities tend to reverse in the subpallial

LSc (Kruskal-Wallis P = 0.06), suggesting that species dif-

ferences in sociality are most closely associated with the

relative densities of binding sites along a dorso-ventral

gradient, as confirmed in (F).


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Peripheral and central the same?















Fig. 4. [I125]-OVTA binding sites in the LS differentiate

territorial and flocking finch species. Hp indicates hippo-

campus; LSc.d, dorsal zone of the caudal lateral septum;

LSc.v,vl, ventral and ventrolateral zones of the caudal

lateral septum; N, nidopallium; PLH, posterolateral hypo-

thalamus; and TeO, optic tectum. (A to C) Representative

autoradiograms of binding sites in two sympatric con-

geners, the territorial violet-eared waxbill (A) and the

gregarious Angolan blue waxbill (B) plus the highly gre-

garious zebra finch (C). (D) Densities of [I125]-OVTA binding

sites in the dorsal (pallial) LSc of two territorial species

(melba finch, MF, and violet-eared waxbill, VEW), a mod-

erately gregarious species (Angolan blue waxbill, ABW),

and two highly gregarious species (spice finch, SF, and

zebra finch, ZF). Territorial and flocking species are shown

in blue and orange, respectively. No sex differences are

observed and sexes were pooled. Total n = 23 (SOM text).

Different letters above the boxes denote significant species

differences (Mann-Whitney P < 0.05) following significant

Kruskal-Wallis. Box plots show the median (line), 75th and

25th percentiles (box), and 95% confidence interval (error

bars). (E) Binding densities tend to reverse in the subpallial

LSc (Kruskal-Wallis P = 0.06), suggesting that species dif-

ferences in sociality are most closely associated with the

relative densities of binding sites along a dorso-ventral

gradient, as confirmed in (F).


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Males?

Oxytocin


Sociality

They say:

I say:Oxytocin

?

Sociality (?)


Chapter 5: A modest proposal

A way to extend this experiment...

Oxytocin


Sociality

Oxytocin

Sociality

What is “sociality”?

- “Species typical group size”?- Preference for familiar conspecifics? (Affiliation)- Preference for large groups? (Flocking)

Here’s my idea:

Let’s put zebra finches in a room,

and let them show us what they

prefer when we give them doses

of mesotocin.

How do we measure their preference?

1

2

4 5

6 7

3 89

1011

How do we measure their preference?Behaviour Level of organization Structure

Affiliation

Group size

Dyadic

Individual / group

How do we measure their preference?

1

2

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6 7

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Aviary

Video

M

M

M

M

M

S

S

S

S

S

M = Mesotocin

S = Saline

O = Oxytocin antagonist

Five conditions:

All mesotocin

M M M M MM M M M M

Half mesotocin

M M M M M

All saline Half antagonist

All antagonist

O O O O OO O O O OS S S S S

S S S S SS S S S S

O O O O OS S S S S

M

M

M

M M

M

Affiliation: dyads pair off preferentially

M

M

M

M

M

S

SS

S

Flocking: mesotocin promotes preferences for larger groups

M

M

S

S

S

S

S

S

S

S

Something else: mesotocin members become important to group structure.

Thank you forlistening!