Population Densy ad Soca athoog When a populatio of borory ras S allowed o crease coned space, he rs evelop acely aoma paters o ehavior th ca eve lead o the exnto he pouao b Jhn n
I
the ceebrated thesis o homas Maths, vice ad misery impose the timate atra imit o the growth o popatios. Stdets o the sbject have give most o their attetio to misery, that is, to predatio, disease ad food sppy as forces that operate to adjst the size o a popatio to its e vironmet. Bt what of vice? Settig side the mora brde of this word, what are the eects o the socia behavior o a species o popatio growthad o popatio desity o soci behavior Some years ago I attempted to sbmit this qestio to experimeta iqiry. coed a popatio o wid Norway rats i a qarteracre ecosre. ith a abdace of food ad pacs to ive ad with predatio ad disease eimi ated or miimized, oy the aims behavior with respect to oe aother re maied as a actor that might aect the icrease i their mber. There cod be o escape rom the behaviora cose qeces o rising popatio desity. By the ed of 27 mths the popatio had become stabiized at 150 adts. Yet adt mortaity was so ow that 5,000 adts might have bee exected from the observed reprodctive rate. The rea so this arger popatio did ot mate riaize was that iat mortaity was ex tremey high. Eve with oy 150 adts i the ecosre, stress rom socia iteracti ed to such disrtio o atera behavior that ew youg srvived. With this backgrod i ind tred to observatio o a domesticated abio strai of the Norway rat nder more cotroed circmstances doors. he data or the reset discssio come rom the histories o six diret popatios. Each was permittd to icrease to approximatey twice the mber that my experiece had idicated cod occpy the avaiabe spae with oy moderate stress rom socia iteractio. I ach
case my associates ad I maitied cose srveiace of the cooies or 6 moths i order to otai detaied records o the modictios o behavior idced by opatio opatio desity he coseqeces o the behaviora pthoogy we observed were most apprt amog the mas. May were nbe to carry pregac to term or to srvive deivery o their itters if they did. A eve greater mber, ater sc cssy givig birth, fe short i their mtera fnctios. Amog the maes the behavior distrbaces raged rom sexa deviatio to caibaism ad from reetic overactivity to a pathoogi ca withdrawa ro which idividas wod emerge to eat, drink ad move abot oy whe other members o the commity were asee. he socia or gnizatio of the aimas showed eqa disrptio. Each o the experimeta popatios divided itse ito severa grops, i each o which the sx ratios were drasticay modied. Oe grop might cosist o six or seve femaes ad oe mae, whereas aother wod have 20 maes ad ony 10 maes. he commo sorce o these distrb aces became most dramaticay appar et i the popatios o or rst series o thre exprimets, i which we ob srvd the deeopmet o what we caed a behaviora sink. he animas wod crowd together i greatest mbe i oe o the or itercoectig pes i which the cooy was maitaied. As ma as 60 o the 80 rats i each ex prieta popatio wod ssembe i oe e drig periods of feedig. Idi viua rats wod rarey at excpt i the comp o other rats. As a rest ex treme uatio desities deveoped i the e adopted or eatig, eaving the others with sparse popuatios. Eatig ad othr bioogica activities wr thereby trnsfored into soci ac
tivities i which the pricipa satisfac tio was iteractio with other rats. I the case of eatig, this trasormatio of behavior did ot keep the aimas rom secrig adeqate tritio. But the same pathoogica "togetheress eded to disrpt the ordered seqeces o activity ivoved i other vita modes of behavior sch as the cortig o sex parters, the bidig o ests ad the rsig d care o the yog. I the experimets i which the behviora sik deveoped, inat mortaity ra as high as 96 per cent amog the most disorieted grops i the popatio. ve i the absece o the behaviora sik, i the secod series o three experimets, iat mortaity reachd 80 per cet amog the correspodig members o the expri meta opatios e desig o the experimets was reativey simpe. he three popa
tios o the rst series each bega with 2 rats; each popatio o the second series bega with 56 rats. I a cases the aimas were jst past weaig ad were evey divided betwee aes ad emaes. B the 12th moth a the popatios had mtipied ad each comprised 80 adts. hereafter remova of the iats that srvived birth ad wea ig hed the popatios stead. A thogh the destrctive eects o popatio desity icreased drig the corse of the experimets, ad the mortaity rate amog the femaes ad amog the yog was mch higher i the 16th moth tha it was earier, the nmber o yog that srvived to weaig was aways arge eogh to oset the eects o adt mortaity ad actay to icrase the popatio The evidece idicates, however, that i time aires o reprodctive c tio wod have cased the cooies to die ot. At the ed of the rst series o exerimets eight ratsthe or heathi
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EFFECT PPLATON ENSITY n te bhir nd sci gniztin f ts s stdid b nning gs f 80 nims in 10·b·1·ft m diidd int ns b n eetid fn A ns (nmbed 3 nd cise m d} mp ding ns Cnic bjs e d ps; trs it t bts dinng t Ed b chd
b winding stics c d nst bxs sn i pn 1 tp f b s bn md Rmps cnncd pns bt 1 nd Rts trf tndd t cncntt in pns nd 3 De mnt f "bi sin ic ft insd pp tin in n n is cted in pn h th ts e ti simtns R ppchng mp i n 3 is n et fm
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iasd hse hsica arre, he sies of the grops i eac o have ee exected o rage from s fe as 13 o as a as 27. Wih the passae of ti, however, chages i ehavior teded o skew the distritio of e rats amog the es eve ore. Of the 100 distic sleeig gros coted i h 10th to 12h moth of each exeri , ol 37 fe withi he expected se rage. I 33 gros there were few er tha 13 ras, ad i 30 gros he o xceded 27 The sex ratio aproxi ated eqalit ol i hose grops tha fell wihi the exected sie rage. he saller gros, geerall composed of ight adlts, her re seldo mor
ha wo males. I h lrger rop, he other had, hre were ma mor males tha feales. As igh e x peced, h smaler grops esalished heselves i the ed pes, whras h larger grops were sall oservd o for i the iddle ps. The fa emers of th polaio disried heselves ao eqall i he for es, t he ale poplatio as co craed almos overwhelmig i the iddle pes. Oe major facor i the creaio of this sae of aairs was he rggle for sats hat oo plae aog th ales. Shorl afer ae ras reach marit a ao six moh of ae, h er io
rod ri of ghs ha va xes heir ositio i he soil rr or exerims sch g a aog h ales i al he pes, o middle ad ed. I he ed es, ho ever, i ecame possile for a sige doia ae o ae over he area as his rrior. Drig he period whe th socia hierarh was i esaished, he sordiae males i pes adoed the ai of arisig ear. his ead hem to ea d drik i pec. Sice ras geera ea i he corse f heir oral waderig, he sori ae rsids of he d pe er i o feed i oe of m s. Wh, after feedig, h ae o
OOD HOPPER used in rs series of eperimes is seen a e f i this drai Waer ra is a te ri Te opper overe ith ire grati ad o ad peles o food, made eain a hy ativiy dui h oe ra as likey mee anoer
us i oserd e deveopme of a beaviora sik: e animas uld ea oy i te presene of oers, an e prefeed ne o e four oppers in te room o a e oers im per en f e animals rodd io e p ontaiig his opper o a
WATER ONTAN used i seond series o eperimens i s a he riht i his drain. ood oppe is a e ef Te founai as opad by essin a . Thus i made rikin a lthy tivity, assoiad it he pesee of ohers Bu i
did no rae a bhaviora sik Ahou e rats oud drik nl if or aimals e ps, hey eaed i is atiity in eir ome pes immedia aftr aakei. The foutai erefoe ad o podue a ev distributio of puation
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oung ui nss fo h and rod h fo ha I an siuaion aos ha a oh onsird a dang o h us sh woud i h infans u on a a i and a h in h ouh o a sa a Nohing ou isa hr fo his as uni h ni i ha n ov Ha h inas o in h ood ns suviv h gnan as od aong h as in h i ns w no ow han hos od in h nd ns u a sa nag of hs gnancis inad in iv ihs In h son sis of xins 80 n o h infans o in h i
ahoog ha an foun wii a go h gn of a havioa sink was fosd h arangns a w ad o ing h anias In hs xins h ood consisd of sa ha s ha k n a cicua ho od wi sh In consqunc sasacion o hung r quid a oninuous o asing sva inus h hans ho w good ha hi on ra was aing aoh ou oin i a h ho As was niond ai as usa a in i houghou hi waking hous hnv h a hung an foo is vaia Sin h aangn o ras o as ino h id pns han ino h os i as i
ns id fo aning I h rs sis 96 n ish fo his i h as i h i ns no ss ac han h as h ssus o ouaion dnsi In oh sis o xins h soia ahoog aong h as as high. In h s sis hov i was o aggva han i was in h son his inas in disuanc ong h n ouans of h s sis o xins was di ad o h dvon of h hnonon of h havioa snh ouo o an havioa oss ha os ani as ogh in unusua ga nus h unhah onnoaions of h a no accina: a havioa sink dos ac o aggava a fs of
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EAVIORAL SINK developed in the rst series of three experimets drain half the rats either into pen or pen 3 o eah room to drin d sleep, ad even more ito that pe to eat Chrt
eribes the situation in the 13th mot of th experiment By then the poplation distributions ere fairl stae and many emales in the dese opuated pens hd died_ Oe mle i room
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OULATON DISTRIBTONS in the seond series of three experimnts, in whi no bevioral sin developd ere more eve than they ere in the rst series and the det rte mo
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emles and infnts as loer hart shos the situatio i the 13th month hen oe male had established pens 3 and of room s is terrtory and nother s taing ovr pen thus
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mn v u su o sus a wi u amon os a sisd a lau a dvod In owdd mdd ns no on in dividul oud o osion n is ia mann In v oup of 12 o mo mas on was mos ssiv an mos on vio in s Nvlss s was odal ousd om s osiion A ula invals duin ous o i wakin ous oankn mals n ad n ffoas umnad n ns o dominan fom on ma o no. In wn s umuu ous anins o uad aiv m pvaild. ssv dominan animas w mos noma mas i ou populaons sdom od fmals o juvnils Y vn xd oasiona sns of aol o oin sk aakin fmals juvnis and lss aiv mals n sowin aua pdilion wi as do no nomall disla
fo in o anmals o il Blow domnan mas o on us sa and n i of vi w omosxuasa ou as dsd as ansxual s animas anl oud no dsmina wn aoa and naoa sx ans mad sxual advans o mas juvnils and mas a w no in sus mas in ludn domans as wll as os o ansxuals' own ou usu al ad anons na vl o aivi o s anima as onl mod w fqu a akd i dominan assoas u v a onndd fo saus wo o s o mal md o o wi ad sind ni om sul fo domnan w owv a xa opos ols as fa s i lvs of aiv w on nd s w omll assiv and movd ou ommun lk somnamuliss inod al o as of o sxs and a
o as od m Evn n mas w sus s assi animas mad no advans o m And onl v al did o mals aak m o aoa m o an kind o a o sual os assiv anmals woud av aad o ais and mos aai mms o h mmuni. w a and slk and i fu sowd non o aks and a sos n in wi mals usual na Bu i soial disonaio was na l om
p a as ss all s md amon mals was ou I av ald s s anmas wi alwas ivd i mdd ns ook no a a al in saus sul Nvlss w mos av of al mas in ximna oulaons and ssd in i aiv in s o aaks domnan anmals In addiion o in paiv ps w o
NORMAL MATERAL BEHAVOR among rats inludes buiding , wellshped nest for the oung. The ding at the left sows suh a nes holding a reent orn itter. The drawing at te rigt shows ths ame nest out wo wees late. It has been
attened the eght of the animals' odes t it sti oes mple poeon and wh n the emaining pups an ti rest omora In these expeimets ha he osping of o ma mothes survive infa nd were suesfu weaned.
ABORMAL MATERNAL EAVIOR shown b females ex· poe to he pessues o popuaton desi inludes aiure to id adequae nets. The dg a e et shows he reen on oung o disturbed femle. She sted to mae net but
ever ished t. Te dawing a he ig shows he oung about two wees lae. One pup has alead eft and anothe is eaving. Nehe an suvive one. In hese expeiments te moait rte among nfns of disubed mothers was as high 96 pe ent.
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hyersexu hsexu a i tie ay f te beae anniba st They ere ys the aert fr estrus fees If tere were e i their n es hey wu ie i wait fr g eris at the ts f te ras that gve the brd es a eer dwn it the. They ways tured ad e s s as the territria rt aught sight f the Eve if they i t age t ese uhurt they wud s retu t their vatage it. Te rbers duted their ursuit f estrus fees i bra a er Mtig ag rats usuay invves distit urtshi ritu I the rst hase f tis riu the e ursues the fee She thereu retires fr a whie it te bur d e e ies quiety it utside siay king his head it the bur fr ent but never eterig it ( I te id frs f te Nrwy rat ths hse usuy ivves a urtshi de n the un t te ut f te burw) The feae t st eerges fr the burrw d a ets the as ves Eve i te disrdered uity f the idde es this tte as bserved by a the es wh egaged i nra etersex u beavir ut the rbers wud nt terte eve a srt erid f waitig at the burrws i te es here aessi be feaes ived As s as feae retired t a burrw rber wud f w er iside On these exeditins the rbers fte fud ded yung ying i the ests s resut they tended t ee aibaisti i te ter nths f a uatins istry
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thugh the bevira sik did t eve i te se series f exeriets te thg exibited by te uatis i bth sets f exeri es ad in es was severe Even i the brd es feaes u raise ny haf teir yug t weaing Nr des te dierene in it rtaity betwee the dde ens f te rst a sed series96 er et i the rst as sed t 80 er et in te sed rereset a bigiay sigiant irveet It is bvius tat the be· havira reertry wit whi the Nrway rt s eerge fr te trias evuti ad estiati ust break dw uder te si ressures generated by uatin desity In tie reeet f exerieta reures and f the iterretatin f these studies ay advne ur understadig t the it where the y ntribute t the akig f vaue judgents abut anagus rbes frnting the hua sees
1,0 The mst siulng cae for sece in Amra tdy n New York S.
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