The Population Ecology of Organizations

STOR

Michael T. Hannan, John Freeman American Journal of Sociology, Volume 82, Issue 5 (Mar., 1977),929-9 !. "#a$le %&'( h##)(**lin+s.,s#or.or-*sici.sici/0002-9 02128197702129821235124929123T56771262.0.471282-J

9our use o: #he J"T7& archi;e in<ica#es =our acce)#ance o: J"T7&> s Terms an< 4on<i#ions o: %se, a;aila$le a# h##)(**???.,s#or.or-*a$ou#i#erms.h#ml. J"T7&>s Terms an< 4on<i#ions o: %se )ro;i<es, in )ar#, #ha# unless =ou ha;e o$#aine< )rior )ermission, =ou ma= no# <o?nloa< an en#ire issue o: a ,ournal or mul#i)le co)ies o: ar#icles, an< =ou ma= use con#en# in #he J"T7& archi;e onl= :or =our )ersonal, non-commercial use. 6ach co)= o: an= )ar# o: a J"T7& #ransmission mus# con#ain #he same co)=ri-h# no#ice #ha# a))ears on #he screen or )rin#e< )a-e o: such #ransmission. American Journal of Sociology is )u$lishe< $= The %ni;ersi#= o: 4hica-o 5ress. 5lease con#ac# #he )u$lisher :or :ur#her )ermissions re-ar<in- #he use o: #his ?or+. 5u$lisher con#ac# in:orma#ion ma= $e o$#aine< a# h##)(**??? .,s#or.or-*,ournals*uc)ress.h#ml.

American Journal of Sociology @1977 The %ni;ersi#= o: 4hica-o 5ress J"T7& an< #he J"T7& lo-o are #ra<emar+s o: J"T7&, an< are &e-is#ere< in #he %.". 5a#en# an< Tra<emar+ 7::ice. For more in:orma#ion on J"T7& con#ac#,s#or-in:oAumich.e<u. @2002J"T7&

h##)(**??? .,s#or.or-* Fri Jan 25 18(22(28 2002

The Population
Michael T. Hannan Stanford University

Ecology of 7r-aniBa#ions>

John Freeman University of California, Berkeley

3 )o)ula#ion ecolo-= )ers)ec#i;e on or-aniBa#ion-en;ironmen# rela #ions is )ro)ose< as an al#erna#i;e #o #he <ominan# a<a)#a#ion )er s)ec#i;e. The s#ren-#h o: iner#ial )ressures on or-aniBa#ional s#ruc#ure su--es#s #he a))lica#ion o: mo<els #ha# <e)en< on com)e#i#ion an< selec#ion in )o)ula#ions o: or-aniBa#ions. "e;eral such mo<els as ?ell as issues #ha# arise in a##em)#s #o a))l= #hem #o #he or-aniBa#ion en;ironmen# )ro$lem are <iscusse<.
I. ICT&7D%4TI7C

3nal=sis o: #he e::ec#s o: en;ironmen# on or-aniBa#ional s#ruc#ure has mo;e< #o a cen#ral )lace in or-aniBa#ions #heor= an< research in recen# =ears. This shi:# has o)ene< a num$er o: eEci#in- )ossi$ili#ies. 3s =e# no#h in- li+e #he :ull )romise o: #he shi:# has $een realiBe<. Fe $elie;e #ha# #he lac+ o: <e;elo)men# is <ue in )ar# #o a :ailure #o $rin- ecolo-ical mo<els #o $ear on Gues#ions #ha# are )reeminen#l= ecolo-ical. Fe ar-ue :or a re:ormula#ion o: #he )ro$lem in )o)ula#ion ecolo-= #erms. 3l#hou-h #here is a ?i<e ;arie#= o: ecolo-ical )ers)ec#i;es, #he= all :ocus on selec#ion. Tha# is, #he= a##ri$u#e )a##erns in na#ure #o #he ac#ion o: selec#ion )rocesses. The $ul+ o: #he li#era#ure on or-aniBa#ions su$scri$es #o a <i::eren# ;ie?, ?hich ?e call #he a<a)#a#ion )es)ec#i;e.5 3ccor<in1This research ?as su))or#e< in )ar# $= -ran#s :rom #he Ca#ional "cience Foun <a#ion (H"-220 5) an< #he ")encer Foun<a#ion. Hel):ul commen#s ?ere )ro;i<e< $= 3mos Ha?le=, Francois Cielsen, John Me=er, Marshall Me=er, Je::re= 5:e::er, an< Ho?ar< 3l<rich. 2 There is a su$#le rela#ionshi) $e#?een selec#ion an< a<a)#a#ion. 3<a)#i;e learn in- :or in<i;i<uals usuall= consis#s o: selec#ion amon- $eha;ioral res)onses. 3<a)#a #ion :or a )o)ula#ion in;ol;es selec#ion amon- #=)es o: mem$ers. More -enerall=, )rocesses in;ol;in- selec#ion can usuall= $e recas# a# a hi-her le;el o: anal=sis as a<a)#a#ion )rocesses. Ho?e;er, once #he uni# o: anal=sis is chosen #here is no am $i-ui#= in <is#in-uishin- selec#ion :rom a<a)#a#ion. 7r-aniBa#ions o:#en a<a)# #o en;ironmen#al con<i#ions in concer# an< #his su--es#s a s=s#ems e::ec#. Thou-h :e? #heoris#s ?oul< <en= #he eEis#ence o: such s=s#ems e::ec#s, mos# <o no# ma+e #hem a su$,ec# o: cen#ral concern. It is im)or#an# #o no#ice #ha#, :rom #he )oin# o: ;ie? em$race< $= sociolo-is#s ?hose in#eres#s :ocus on #he $roa<er social s=s#em, selec#ion in :a;or o: or-aniBa#ions ?i#h one se# o: )ro)er#ies #o #he <is:a;or o: #hose ?i#h o#hers is o:#en an a<a)#i;e )rocess. "ocie#ies an< communi#ies ?hich consis# in )ar# o: :ormal or-aniBa#ions a<a)# )ar#l= #hrou-h )rocesses #ha# a<,us# #he miE#ure o:

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#o #he a<a)#a#ion )ers)ec#i;e, su$uni#s o: #he or-aniBa#ion, usuaII= man a-ers or <ominan# coali#ions, scan #he rele;an# en;ironmen# :or o))or#u ni#ies an< #hrea#s, :ormula#e s#ra#e-ic res)onses, an< a<,us# or-aniBa#ional s#ruc#ure a))ro)ria#el=. The a<a)#a#ion )ers)ec#i;e is seen mos# clearl= in #he li#era#ure on mana-emen#. 4on#ri$u#ors #o i# usuall= assume a hierarch= o: au#hori#= an< con#rol #ha# loca#es <ecisions concernin- #he or-ania#ion as a ?hole a# #he #o). It :oIIo?s, #hen, #ha# or-aniBa#ions are a::ec#e< $=

#heir en;i ronmen#s accor<in- #o #he ?a=s in ?hich mana-ers or lea<ers :ormula#e s#ra#e-ies, ma+e <ecisions, an< im)lemen# #hem. 5ar#icularl= success:ul mana-ers are a$le ei#her #o $u::er #heir or-aniBa#ions :rom en;ironmen#al <is#ur$ances or #o arran-e smoo#h a<,us#men#s #ha# reGuire minimal <is ru)#ion o: or-aniBa#ional s#ruc#ure. 3 similar )ers)ec#i;e, o:#en ?or<e< <i::eren#l=, <omina#es #he socio lo-ical li#era#ure on #he su$,ec#. It )la=s a cen#ral role in 5arsons>s (195 ) :unc#ional anal=sis o: or-aniBa#ion-en;ironmen# rela#ions an< i# is :oun< in #he more s#ric#l= Fe$erian #ra<i#ion (see "elBnic+ 1957). It is in#er es#in- #o no#e #ha#, ?hile :unc#ionalis#s ha;e $een in#eres#e< in s=s#em e::ec#s an< ha;e $ase< much o: #he lo-ic o: #heir a))roach on sur;i;al im)era#i;es, #he= ha;e no# <eal# ?i#h selec#ion )henomena. This is )ro$ a$l= a reac#ion a-ains# or-aniBa#ion #heor= ?hich re:lec#s social Dar?in ism. 6Echan-e #heoris#s ha;e also em$race< #he a<a)#a#ion )ers)ec#i;e ('e ;ine an< Fhi#e 19 1). 3n< i# is na#ural #ha# #heories em)hasiBin- <eci sion ma+in- #a+e #he a<a)#a#ion ;ie? (March an< "imon 1958I 4=er# an< March 19 2). 6;en Thom)son>s (19 7) cele$ra#e< marria-e o: o)en-s=s#ems an< close<-s=s#ems #hin+in- em$race< #he a<a)#a#ion )er s)ec#i;e eE)lici#l= (see )ar#icularl= #he secon< hal: o: Thom)son>s $oo+). 4learl=, lea<ers o: or-aniBa#ions <o :ormula#e s#ra#e-ies an< or-aniBa #ions <o a<a)# #o en;ironmen#al con#in-encies. 3s a resul# a# leas# some o: #he rela#ionshi) $e#?een s#ruc#ure an< en;ironmen# mus# re:lec# a<a) #i;e $eha;ior or learnin-. 8u# #here is no reason #o )resume #ha# #he -rea# s#ruc#ural ;aria$ili#= amon- or-aniBa#ions re:lec#s onl= or e;en )rimaril= a<a)#a#ion. There are a num$er o: o$;ious limi#a#ions on #he a$ili#= o: or-aniBa #ions #o a<a)#. Tha# is, #here are a num$er o: )rocesses #ha# -enera#e s#ruc#ural iner#ia. The s#ron-er #he )ressures, #he lo?er #he or-aniBa#ions>
;arious +in<s o: or-aniBa#ions :oun< ?i#hin #hem. Fhereas a com)le#e #heor= o: or-aniBa#ion an< en;ironmen# ?oul< ha;e #o consi<er $o#h a<a)#a#ion an< selec#ion, reco-niBin- #ha# #he= are com)lemen#ar= )rocesses, our )ur)ose here is #o sho? ?ha# can $e learne< :rom s#u<=in- selec#ion alone (see 3l<rich an< 5:e::er J197 K :or a s=n#he#ic re;ie? o: #he li#era#ure :ocusin- on #hese <i::eren# )ers)ec#i;es).

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a<a)#i;e :leEi$ili#= an< #he more li+el= #ha# #he lo-ic o: en;ironmen#al se lec#ion is a))ro)ria#e. 3s a conseGuence, #he issue o: s#ruc#ural iner#ia is cen#ral #o #he choice $e#?een a<a)#a#ion an< selec#ion mo<els. The )ossi$ili#= #ha# or-aniBa#ion s#ruc#ure con#ains a lar-e iner#ial com)onen# ?as su--es#e< $= 8urns an< "#al+er (19 1) an< "#inchcom$e (19 5). 8u#, on #he ?hole #he su$,ec# has $een i-nore<. 3 num$er o: rele;an# )ro)osi#ions can $e :oun< in #he or-aniBa#ions li#era#ure, ho?e;er. Iner#ial )ressures arise :rom $o#h in#ernal s#ruc#ural arran-emen#s an< en;ironmen#al cons#rain#s. 3 minimal lis# o: #he cons#rain#s arisin- :rom in#ernal consi<era#ions :ollo?s. 1. 3n or-aniBa#ion>s in;es#men# in )lan#, )ersonnel cons#i#u#es asse#s #ha# are no# easil= or :unc#ions. The ?a=s in ?hich such sun+ o)#ions are so o$;ious #ha# #he= nee< no# $e eGui)men#, an< s)ecialiBe< #rans:era$le #o o#her #as+s cos#s cons#rain a<a)#a#ion <iscusse< :ur#her.

2. 7r-aniBa#ional <ecision ma+ers also :ace cons#rain#s on #he in:orma #ion #he= recei;e. Much o: ?ha# ?e +no? a$ou# #he :lo? o: in:orma#ion #hrou-h or-aniBa#ional s#ruc#ures #ells us #ha# lea<ers <o no# o$#ain an= #hin- close #o :ull in:orma#ion on ac#i;i#ies ?i#hin #he or-aniBa#ion an< en;ironmen#al con#in-encies :acin- #he su$uni#s. 2. In#ernal )oli#ical cons#rain#s are e;en more im)or#an#. Fhen or-a niBa#ions al#er s#ruc#ure, )oli#ical eGuili$ria are <is#ur$e<. 3s lon- as #he )ool o: resources is :iEe<, s#ruc#ural chan-e almos# al?a=s in;ol;es re<is #ri$u#ion o: resources across su$uni#s. "uch re<is#ri$u#ion u)se#s #he )re ;ailin- s=s#em o: eEchan-e amon- su$uni#s (or su$uni# lea<ers). "o a# leas# some su$uni#s are li+el= #o resis# an= )ro)ose< reor-aniBa#ion. More o;er, #he $ene:i#s o: s#ruc#ural reor-aniBa#ion are li+el= #o $e $o#h -en eraliBe< (<esi-ne< #o $ene:i# #he or-aniBa#ion as a ?hole) an< lon--run. 3n= ne-a#i;e )oli#ical res)onse ?ill #en< #o -enera#e shor#-run cos#s #ha# are hi-h enou-h #ha# or-aniBa#ional lea<ers ?ill :ore-o #he )lanne< reor -aniBa#ion. (For a more eE#ensi;e <iscussion o: #he ?a=s in ?hich #he in#ernal )oli#ical econom= o: or-aniBa#ions im)e<es chan-e or a<a)#a#ion, see Do?ns J19 7J an< Lal< J1970J.) !. Finall=, or-aniBa#ions :ace cons#rain#s -enera#e< $= #heir o?n his #or=. 7nce s#an<ar<s o: )roce<ure an< #he alloca#ion o: #as+s an< au #hori#= ha;e $ecome #he su$,ec# o: norma#i;e a-reemen#, #he cos#s o: chan-e are -rea#l= increase<. Corma#i;e a-reemen#s cons#rain a<a)#a#ion in a# leas# #?o ?a=s. Firs#, #he= )ro;i<e a ,us#i:ica#ion an< an or-aniBin)rinci)le :or #hose elemen#s #ha# ?ish #o resis# reor-aniBa#ion (i.e., #he= can resis# in #erms o: a share< )rinci)le). "econ<, norma#i;e a-reemen#s )reclu<e #he serious consi<era#ion o: man= al#erna#i;e res)onses. For eE am)le, :e? research-orien#e< uni;ersi#ies seriousl= consi<er a<a)#in- #o

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<eclinin- enrollmen#s $= elimina#in- #he #eachin- :unc#ion. To en#er#ain #his o)#ion ?oul< $e #o challen-e cen#ral or-aniBa#ional norms.M The eE#ernal )ressures #o?ar< iner#ia seem #o $e a# leas# as s#ron-. The= inclu<e a# leas# #he :ollo?in- :ac#ors. 1. 'e-al an< :iscal $arriers #o en#r= an< eEi# :rom mar+e#s ($roa<l= <e:ine<) are numerous. Discussions o: or-aniBa#ional $eha;ior #=)icall= em)hasiBe $arriers #o en#r= (s#a#e license< mono)ol= )osi#ions, e#c.). 8arriers #o eEi# are eGuall= in#eres#in-. There are an increasin- num$er o: ins#ances in ?hich )oli#ical <ecisions )re;en# :irms :rom a$an<onincer#ain ac#i;i#ies. 3ll such cons#rain#s on en#r= an< eEi# limi# #he $rea<#h o: a<a)#a#ion )ossi$ili#ies. 2. In#ernal cons#rain#s u)on #he a;aila$ili#= o: in:orma#ion are )aral lele< $= eE#ernal cons#rain#s. The acGuisi#ion o: in:orma#ion a$ou# rele ;an# en;ironmen#s is cos#l= )ar#icularl= in #ur$ulen# si#ua#ions ?here #he in:orma#ion is mos# essen#ial. In a<<i#ion, #he #=)e o: s)ecialis#s em)lo=e< $= #he or-aniBa#ion cons#rains $o#h #he na#ure o: #he in:orma#ion i# is li+el= #o o$#ain (see Hrano;e##er 1972) an< #he +in< o: s)ecialiBe< in:or ma#ion i# can )rocess an< u#iliBe. 2. 'e-i#imac= cons#rain#s also emana#e :rom #he en;ironmen#. 3n= le-i#imac= an or-aniBa#ion has $een a$le #o -enera#e cons#i#u#es an asse# in mani)ula#in- #he en;ironmen#. To #he eE#en# #ha# a<a)#a#ion (e.-., elimina#in- un<er-ra<ua#e ins#ruc#ion in )u$lic uni;ersi#ies) ;iola#es #he le-i#imac= claims, i# incurs consi<era$le cos#s. "o eE#ernal le-i#imac= con si<era#ions also #en< #o limi# a<a)#a#ion. !. Finall=, #here is #he collec#i;e ra#ionali#= )ro$lem. 7ne o: #he mos# <i::icul# issues in con#em)orar= economics concerns -eneral eGuili$ria. If

one can :in< an o)#imal s#ra#e-= :or some in<i;i<ual $u=er or seller in a com)e#i#i;e mar+e#, it <oes no# necessaril= :ollo? #ha# #here is a -eneral eGuili$rium once all )la=ers s#ar# #ra<in-. More -enerall=, i# is <i::icul# #o es#a$lish #ha# a s#ra#e-= #ha# is ra#ional :or a sin-le <ecision ma+er ?ill $e ra#ional if a<o)#e< $= a lar-e num$er o: <ecision ma+ers. 3 num$er o: solu#ions #o #his )ro$lem ha;e $een )ro)ose< in com)e#i#i;e mar+e# #heor=, $u# ?e +no? o: no #rea#men# o: #he )ro$lem :or or-aniBa#ions -enerall=. %n#il such a #rea#men# is es#a$lishe< ?e shoul< no# )resume #ha# a course o: ac#ion #ha# is a<a)#i;e :or a sin-le or-aniBa#ion :acinsome chan-in- en;ironmen# ?ill $e a<a)#i;e :or man= com)e#in- or-ani Ba#ions a<o)#in- a similar s#ra#e-=. 3 num$er o: #hese iner#ial )ressures can $e accommo<a#e< ?i#hin #he a<a)#a#ion :rame?or+. Tha# is, one can mo<i:= an< limi# #he )ers)ec#i;e in or<er #o consi<er choices ?i#hin #he cons#raine< se# o: al#erna#i;es. 8u#
2 Me=er>s (1970) <iscussion o: an or-aniBa#ion>s char#er a<<s :ur#her su))or# #o #he ar-umen# #ha# norma#i;e a-reemen#s arri;e< a# earl= in an or-aniBa#ion>s his#or= cons#rain -rea#l= #he or-aniBa#ion>s ran-e o: a<a)#a#ion #o en;ironmen#al cons#rain#s.

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#o <o so -rea#l= limi#s #he sco)e o: one>s in;es#i-a#ion. Fe ar-ue #ha# in or<er #o <eal ?i#h #he ;arious iner#ial )ressures #he a<a)#a#ion )ers)ec#i;e mus# $e su))lemen#e< ?i#h a selec#ion orien#a#ion. Fe consi<er :irs# #?o $roa< issues #ha# are )reliminar= #o ecolo-ical mo<elin-. The :irs# concerns a))ro)ria#e uni#s o: anal=sis. T=)ical anal =ses o: #he rela#ion o: or-aniBa#ions #o en;ironmen#s #a+e #he )oin# o: ;ie? o: a sin-le or-aniBa#ion :acin- an en;ironmen#. Fe ar-ue :or an eE )lici# :ocus on )o)ula#ions o: or-aniBa#ions. The secon< $roa< issue con cerns #he a))lica$ili#= o: )o)ula#ion ecolo-= mo<els #o #he s#u<= o: human social or-aniBa#ion. 7ur su$s#an#i;e )ro)osal $e-ins ?i#h Ha?le=>s (195.0, 19 8) classic s#a#emen# on human ecolo-=. Fe see+ #o eE#en< Ha?le=>s ?or+ in #?o ?a=s( $= usin- eE)lici# com)e#i#ion mo<els #o s)eci:= #he )rocess )ro<ucin- isomor)hism $e#?een or-aniBa#ional s#ruc#ure an< en ;ironmen#al <eman<s, an< $= usin- niche #heor= #o eE#en< #he )ro$lem #o <=namic en;ironmen#s. Fe ar-ue #ha# Ha?le=>s )ers)ec#i;e, mo<i:ie< an< eE#en<e< in #hese ?a=s, ser;es as a use:ul s#ar#in- )oin# :or )o)ula #ion ecolo-= #heories o: or-aniBa#ions.

II.

575%'3TI7C THICNICH IC TH6 "T%D9 7F 7&H3CIL3TI7C 6CVI&7CM6CT &6'3TI7C"

'i##le a##en#ion is )ai< in #he or-aniBa#ions li#era#ure #o issues concernin)ro)er uni#s o: anal=sis (Freeman 1975). In :ac#, choice o: uni# is #rea#e< so casuall= as #o su--es# #ha# i# is no# an issue. Fe sus)ec# #ha# #he o))o si#e is #rue-#ha# #he choice o: uni# in;ol;es su$#le issues an< has :ar reachin- conseGuences :or research ac#i;i#=. For ins#ance, in #he case a# han<, i# <e#ermines ?hich o: se;eral ecolo-ical li#era#ures can $e $rou-h# #o $ear on #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions. The com)arison o: uni# choice :acin- #he or-aniBa#ional anal=s# ?i#h #ha# :acin- #he $ioecolo-is# is ins#ruc#i;e. To o;ersim)li:= some?ha#, ecolo-ical anal=sis is con<uc#e< a# #hree le;els( in<i;i<ual, )o)ula#ion, an< commu ni#=. 6;en#s a# one le;el almos# al?a=s ha;e conseGuences a# o#her le;els. Des)i#e #his in#er<e)en<ence, )o)ula#ion e;en#s canno# $e re<uce< #o in<i ;i<ual e;en#s (since in<i;i<uals <o no# re:lec# #he :ull -ene#ic ;aria$ili#= o: #he )o)ula#ion) an< communi#= e;en#s canno# $e sim)l= re<uce< #o )o)u la#ion e;en#s. 8o#h #he la##er em)lo= a )o)ula#ion )ers)ec#i;e ?hich is no# a))ro)ria#e a# #he in<i;i<ual le;el. The si#ua#ion :ace< $= #he or-aniBa#ions anal=s# is more com)leE. In s#ea< o: #hree le;els o: anal=sis, he :aces a# leas# :i;e( (I) mem$ers, (2) su$uni#s, (2) in<i;i<ual or-aniBa#ions, (!) )o)ula#ions o: or-aniBa#ions, an< (5) communi#ies o: ()o)ula#ions o:) or-aniBa#ions. 'e;els 2-5 can $e seen as corres)on<in- #o #he #hree le;els <iscusse< :or -eneral ecolo-=, ?i#h #he in<i;i<ual or-aniBa#ion #a+in- #he )lace o: #he in<i;i<ual or-an165

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ism. The a<<e< com)leEi#= arises $ecause or-aniBa#ions are more nearl= <ecom)osa$le in#o cons#i#uen# )ar#s #han are or-anisms. In<i;i<ual mem $ers an< su$uni#s ma= mo;e :rom or-aniBa#ion #o or-aniBa#ion in a man ner ?hich has no )arallel in nonhuman or-aniBa#ion. Ins#ances o: #heor= an< research <ealin- ?i#h #he e::ec#s o: en;iron men#s on or-aniBa#ions are :oun< a# all :i;e le;els. For eEam)le, 4roBier>s ?ell-+no?n anal=sis o: #he e::ec#s o: cul#ure on $ureaucrac= :ocuses on #he cul#ural ma#erials mem$ers $rin- #o or-aniBa#ions (19 !). 3# #he o#her en< o: #he con#inuum ?e :in< anal=ses o: Mor-aniBa#ional :iel<sM (Tur+ 1970I 3l<rich an< &eiss 197 ). 8u#, #he mos# common :ocus is on the or-aniBa#ion an< its en;ironmen#. In :ac#, #his choice is so ?i<es)rea< #ha# #here a))ears #o $e a #aci# un<ers#an<in- #ha# in<i;i<ual or-aniBa #ions are #he a))ro)ria#e uni#s :or #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions. Fe ar-ue :or a )arallel <e;elo)men# o: #heor= an< research a# #he )o)u la#ion (an<, ul#ima#el=, #he communi#=) le;el. 8ecause o: #he <i::erino)inions a$ou# le;els o: anal=sis, M)o)ula#ionM has a# leas# #?o re:eren#s. 4on;en#ional #rea#men#s o: human ecolo-= su--es# #ha# #he )o)ula#ions rele;an# #o #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions are #hose a--re-a#es o: mem$ers a##ache< #o #he or-aniBa#ion or, )erha)s, ser;e< $= #he or-aniBa#ion. In #his sense, #he or-aniBa#ion is ;ie?e< as analo-ue #o a communi#=( i# has collec#i;e means o: a<a)#in- #o en;ironmen#al si#ua#ions. The uni# charac#er o: a )o)ula#ion so <e:ine< <e)en<s on share< :a#e. 3ll mem$ers share #o some eE#en# in #he conseGuences o: or-aniBa#ional success or :ailure. Fe use #he #erm )o)ula#ion in a secon< sense( #o re:er #o a--re-a#es o: or-aniBa#ions ra#her #han mem$ers. 5o)ula#ions o: or-aniBa#ions mus# $e ali+e in some res)ec#, #ha# is, #he= mus# ha;e some uni# charac#er. %n:or#una#el=, i<en#i:=in- a )o)ula#ion o: or-aniBa#ions is no sim)le ma##er. The ecolo-ical a))roach su--es#s #ha# one :ocus on common :a#e ?i#h res)ec# #o en;ironmen#al ;aria#ions. "ince all or-aniBa#ions are <is #inc#i;e, no #?o are a::ec#e< i<en#icall= $= an= -i;en eEo-enous shoc+. Ce;er#heless, ?e can i<en#i:= classes o: or-aniBa#ions ?hich are rela#i;el= homo-eneous in #erms o: en;ironmen#al ;ulnera$ili#=. Co#ice #ha# #he )o)ula#ions o: in#eres# ma= chan-e some?ha# :rom in;es#i-a#ion #o in;es #i-a#ion <e)en<in- on #he anal=s#>s concern. 5o)ula#ions o: or-aniBa#ions re:erre< #o are no# immu#a$le o$,ec#s in na#ure $u# are a$s#rac#ions use:ul :or #heore#ical )ur)oses. If ?e are #o :ollo? #he lea< o: )o)ula#ion $iolo-is#s, ?e mus# i<en#i:= an analo-ue #o #he $iolo-is#>s no#ion o: s)ecies. Various s)ecies are <e :ine< ul#ima#el= in #erms o: -ene#ic s#ruc#ure. 3s Mono< (1971) in<ica#es, i# is use:ul #o #hin+ o: #he -ene#ic con#en# o: an= s)ecies as a $lue)rin#. The $lue)rin# con#ains #he rules :or #rans:ormin- ener-= in#o s#ruc#ure. 1

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4onseGuen#l= all o: #he a<a)#i;e ca)aci#= o: a s)ecies is summariBe< in #he $lue)rin#. If ?e are #o i<en#i:= a s)ecies analo-ue :or or-aniBa#ions,

?e mus# search :or such $lue)rin#s. These ?ill consis# o: rules or )roce <ures :or o$#ainin- an< ac#in- u)on in)u#s in or<er #o )ro<uce an or-ani Ba#ional )ro<uc# or res)onse. The #=)e o: $lue)rin# one i<en#i:ies <e)en<s on su$s#an#i;e concerns. For eEam)le, Marscha+ an< &a<ner (1972) em)lo= #he #erm Mor-aniBa #ional :ormMO #o charac#eriBe #he +e= elemen#s o: #he $lue)rin# as seen ?i#hin a <ecision-ma+in- :rame?or+. For #hem #he $lue)rin# or :orm has #?o :unc#ions( an in:orma#ion :unc#ion #ha# <escri$es #he rules use< in o$#ainin-, )rocessin-, an< #ransmi##in- in:orma#ion a$ou# #he s#a#es o: eE#ernal en;ironmen#s, an< an ac#i;i#= :unc#ion #ha# s#a#es #he rules use< in ac#in- on recei;e< in:orma#ion so as #o )ro<uce an or-aniBa#ional res)onse. To #he eE#en# #ha# one can i<en#i:= classes o: or-aniBa#ions #ha# <i::er ?i#h re-ar< #o #hese #?o :unc#ions, one can es#a$lish classes or :orms o: or-aniBa#ion. "ince our concerns eE#en< $e=on< <ecision ma+in-, ho?e;er, ?e :in< Marscha+ an< &a<ner>s <e:ini#ion o: :orms #oo limi#in-. In :ac#, #here is no reason #o limi# a )riori #he ;arie#= o: rules or :unc#ions #ha# ma= <e:ine rele;an# $lue)rin#s. "o :or us, an or-aniBa#ional :orm is a $lue)rin# :or or-aniBa#ional ac#ion, :or #rans:ormin- in)u#s in#o ou#)u#s. The $lue )rin# can usuall= $e in:erre<, al$ei# in some?ha# <i::eren# ?a=s, $= eEam inin- an= o: #he :ollo?in-( (I) #he :ormal s#ruc#ure o: #he or-aniBa#ion in #he narro? sense-#a$les o: or-aniBa#ion, ?ri##en rules o: o)era#ion, e#c.I (2) #he )a##erns o: ac#i;i#= ?i#hin #he or-aniBa#ion-?ha# ac#uall= -e#s <one $= ?homI or (2) #he norma#i;e or<er-#he ?a=s o: or-aniBin#ha# are <e:ine< as ri-h# an< )ro)er $= $o#h mem$ers an< rele;an# sec #ors o: #he en;ironmen#. To com)le#e #he s)ecies analo-ue, ?e mus# search :or Guali#a#i;e <i: :erences amon- :orms. I# seems mos# li+el= #ha# ?e ?ill :in< such <i::er ences in #he :irs# an< #hir< areas lis#e< a$o;e, :ormal s#ruc#ure an< nor ma#i;e or<er. The la##er o::ers )ar#icularl= in#ri-uin- )ossi$ili#ies. Fhen e;er #he his#or= o: an or-aniBa#ion, i#s )oli#ics, an< i#s social s#ruc#ure are enco<e< in a norma#i;e claim (e.-., )ro:essionaliBa#ion an< colle-ial au#hori#=), one can use #hese claims #o i<en#i:= :orms an< <e:ine )o)u la#ions :or research. Ha;in- <e:ine< #he or-aniBa#ional :orm, ?e can )ro;i<e a more )recise <e:ini#ion o: a )o)ula#ion o: or-aniBa#ions. Jus# as #he or-aniBa#ional an al=s# mus# choose a uni# o: anal=sis, so mus# he choose a s=s#em :or s#u<=. "=s#ems rele;an# #o #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions are
4 The #erm Mor-aniBa#ional :ormM is use< ?i<el= in #he sociolo-ical li#era#ure "#inchcom$e 19 5).

(see

1 7

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6colo-= o: 7r-aniBa#ions

usuall= <e:ine< $= -eo-ra)h=, $= )oli#ical $oun<aries, $= mar+e# or )ro< uc# consi<era#ions, e#c. Hi;en a s=s#ems <e:ini#ion, a )o)ula#ion o: or-a niBa#ions consis#s o: all #he or-aniBa#ions ?i#hin a )ar#icular $oun<ar= #ha# ha;e a common :orm. Tha# is, #he )o)ula#ion is #he :orm as i# eEis#s or is realiBe< ?i#hin a s)eci:ie< s=s#em. 8o#h uses o: #he #erm )o)ula#ion (an< #he ecolo-ical #heories im)lie< #here$=) are li+el= #o )ro;e $ene:icial #o #he s#u<= o: or-aniBa#ional s#ruc #ure. The :irs#, more common, ;ie? su--es#s #ha# or-aniBa#ional s#ruc#ure ou-h# #o $e ;ie?e< as an ou#come o: a collec#i;e a<a)#i;e )rocess. 3ccor< in- #o #his ;ie?, s#ruc#ure an< chan-e ou-h# #o <e)en< on #he a<a)#i;e ness o: su$uni#s an< on #he <i::eren#ial access o: su$uni#s #o en;iron men#al resources. The secon< ;ie? i-nores #he a<a)#i;e ac#i;i#ies o: ele men#s ?i#hin #he or-aniBa#ion eEce)# as #he= cons#i#u#e or-aniBa#ional s#ruc#ure. It :ocuses on #he or-aniBa#ion as an a<a)#in- uni#. 4er#ainl=

$o#h )ers)ec#i;es are nee<e<. Fe are concerne< here onl= ?i#h #he la##er, ho?e;er. Finall=, ?e ?oul< li+e #o i<en#i:= #he )ro)er#ies o: )o)ula#ions mos# in#eres#in- #o )o)ula#ion ecolo-is#s. The main concern in #his re-ar< ?as eE)resse< clearl= $= 6l#on (1927)( MIn sol;in- ecolo-ical )ro$lems ?e are concerne< ?i#h what animals do in #heir ca)aci#= as ?hole, li;inanimals, no# as <ea< animals or as a series o: )ar#s o: animals. Fe ha;e neE# #o s#u<= #he circums#ances un<er ?hich #he= <o #hose #hin-s, an<, mos# im)or#an# o: all, #he limi#in- :ac#ors ?hich )re;en# #hem :rom <oincer#ain o#her #hin-s. 8= sol;in- #hese Gues#ions i# is )ossi$le #o <isco;er #he reasons :or the distribution and numbers of animals in nature." Hu#ch inson (1959) in #he su$#i#le #o his :amous essa=, MHoma-e #o "an#a &osalia,M eE)resse< #he main :ocus e;en more succinc#l=( MFh= 3re There "o Man= Nin<s o: 3nimals.M Ta+in- our lea< :rom #hese <is#in-uishe< ecolo-is#s, ?e su--es# #ha# a )o)ula#ion ecolo-= o: or-aniBa#ions mus# see+ #o un<ers#an< #he <is#ri$u#ions o: or-aniBa#ions across en;ironmen#al con<i#ions an< #he limi#a#ions on or-aniBa#ional s#ruc#ures in <i::eren# en;ironmen#s, an< more -enerall= see+ #o ans?er #he Gues#ion, Fh= are #here so man= +in<s o: or-aniBa#ions.
III. DI"47CTIC%ITI6" IC 647'7HI43' 3C3'9"I"

%#iliBa#ion o: mo<els :rom ecolo-= in #he s#u<= o: or-aniBa#ions )oses a num$er o: anal=#ic challen-es in;ol;in- <i::erences $e#?een human an< nonhuman or-aniBa#ions ?i#h re-ar< #o #heir essen#ial in-re<ien#s. 4on si<er, :irs#, #he non-ene#ic #ransmission o: in:orma#ion. 8iolo-ical anal=ses are -rea#l= sim)li:ie< $= #he :ac# #ha# mos# use:ul in:orma#ion concernina<a)#a#ion #o #he en;ironmen# (?hich in:orma#ion ?e call s#ruc#ure) is #ransmi##e< -ene#icall=. Hene#ic )rocesses are so nearl= in;arian# #ha# 1 8

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eE#reme con#inui#= in s#ruc#ure is #he rule. The small num$er o: im)er :ec#ions -enera#es s#ruc#ural chan-es, ?hich, if acce)#e< $= #he en;iron

men#, ?ill $e #ransmi##e< ?i#h near in;ariance. The eE#reme s#ruc#ural in;ariance o: s)ecies -rea#l= sim)li:ies #he )ro$lem o: <elimi#in- an< i<en#i:=in- )o)ula#ions. More im)or#an#, #he a<a)#i;eness o: s#ruc#ure can $e unam$i-uousl= i<en#i:ie< ?i#h ne# re)ro<uc#ion ra#es. Fhen a )o)ula#ion ?i#h -i;en )ro)er#ies increases i#s ne# re)ro<uc#ion ra#e :ol lo?in- an en;ironmen#al chan-e, i# :ollo?s #ha# i# is $ein- selec#e< :or. This is ?h= mo<ern $iolo-is#s ha;e narro?e< #he <e:ini#ion o: :i#ness #o #he ne# re)ro<uc#i;e ra#e o: )o)ula#ion. Human social or-aniBa#ion )resuma$l= re:lec#s a -rea#er <e-ree o: learnin- or a<a)#a#ion. 3s a resul# i# is more <i::icul# #o <e:ine :i#ness in a )recise ?a=. %n<er a# leas# some con<i#ions, or-aniBa#ions ma= un<er-o such eE#reme s#ruc#ural chan-e #ha# #he= shi:# :rom one :orm #o ano#her. 3s a resul#, eE#reme a<a)#a#ion ma= -i;e rise #o o$ser;e< chan-es #ha# mimic selec#ion. This is )ar#icularl= )ro$lema#ic ?hen #he ;arious or-a niBa#ional :orms are similar on man= <imensions. Fe ha;e ar-ue< )re;iousl= (Hannan an< Freeman 197!) :or a com )osi#e measure o: :i#ness #ha# inclu<es $o#h selec#ion (ac#ual loss o: or-a niBa#ions) an< mo$ili#= amon- :orms (eE#reme a<a)#a#ion). Fi#ness ?oul< #hen $e <e:ine< as #he )ro$a$ili#= #ha# a -i;en :orm o: or-aniBa#ion ?oul< )ersis# in a cer#ain en;ironmen#. Fe con#inue #o $elie;e #ha# such an a))roach has ;alue, $u# ?e no? $elie;e #ha# i# is )rema#ure #o com$ine a<a)#a#ion an< selec#ion )rocesses. The :irs# or<er o: $usiness is #o s#u<= selec#ion )rocesses :or #hose si#ua#ions in ?hich iner#ial )ressures are su: :icien#l= s#ron- #ha# mo$ili#= amon- :orms is unli+el=. Fur#hermore, i# is ?or#h no#in- #ha# #he ca)aci#= #o a<a)# is i#sel: su$,ec# #o e;olu#ion (i.e., #o s=s#ema#ic selec#ion). 3s ?e ar-ue $elo?, or-aniBa#ions <e;elo) #he ca)aci#= #o a<a)# a# #he cos# o: lo?ere< )er :ormance le;els in s#a$le en;ironmen#s. Fhe#her or no# such a<a)#a$le or-aniBa#ional :orms ?ill sur;i;e (i.e., resis# selec#ion) <e)en<s on #he na#ure o: #he en;ironmen# an< #he com)e#i#i;e si#ua#ion. There:ore, a selec#ion )oin# o: ;ie? #rea#s hi-h le;els o: a<a)#a$ili#= as )ar#icular e;olu#ionar= ou#comes. There is a secon< sense in ?hich human ecolo-= a))ears #o <i::er :rom $ioecolo-=. 8lau an< "co## (19 2) )oin# ou# #ha#, unli+e #he usual $io lo-ical si#ua#ion, in<i;i<ual or-aniBa#ions (an< )o)ula#ions o: or-aniBa #ions) ha;e #he )o#en#ial #o eE)an< almos# ?i#hou# limi#. The eE)an< a$ili#= o: )rimi#i;e elemen#s is a )ro$lem $ecause o: our :ocus on #he <is#ri$u#ion o: or-aniBa#ional :orms o;er en;ironmen#s. 3 -i;en :orm (e.-., :ormal $ureaucrac=) can eE)an< #hrou-hou# some s=s#em, mar+e#, or ac#i;i#=, ei#her $ecause one $ureaucrac= -ro?s or $ecause man= $ureau cracies are :oun<e<. 6i#her )rocess ?ill -enera#e an increase in #he )re;1 9

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alence o: $ureaucra#ic or-aniBa#ional ac#i;i#=. 3 li#eral a))lica#ion o: )o) ula#ion ecolo-= #heor= #o #he )ro$lem o: or-aniBa#ional chan-e ?oul< in;ol;e sim)l= coun#in- rela#i;e num$ers in )o)ula#ions. "uch a )roce<ure ma= miss a )henomenon o: cen#ral in#eres# #o #he or-aniBa#ional anal=s#. Fin#er (19 !), in <iscussin- #he anal=#ic )ro$lem raise< here, su--es#s <is

#in-uishin- $e#?een sur;i;al, ?hich <escri$es #he :a#e o: in<i;i<ual or-ani Ba#ions, an< ;ia$ili#=, ?hich <escri$es #he Mshare o: mar+e#M o: a -i;en or-aniBa#ional :orm. Fe :in< a# leas# as much meri# in ano#her )ers)ec#i;e on #he issue o: siBe. Man= #heoris#s ha;e asser#e< #ha# s#ruc#ural chan-e a##en<s -ro?#hI in o#her ?or<s, a sin-le or-aniBa#ion canno# -ro? in<e:ini#el= an< s#ill main#ain i#s ori-inal :orm. For ins#ance, a mouse coul< no# )ossi$l= main #ain #he same )ro)or#ion o: $o<= ?ei-h# #o s+ele#al s#ruc#ure ?hile -ro? in- as $i- as a house. I# ?oul< nei#her loo+ li+e a mouse nor o)era#e )h=siolo-icall= li+e a mouse. 8oul<in- (1952) an< Haire (1959) ar-ue #ha# #he same is #rue :or or-aniBa#ions. 4a)lo? (1957), $uil<in- on ?or+ $= Hraicunas (1922) an< o#hers, ar-ues #ha# #he a$ili#= o: each mem$er o: an or-aniBa#ion #o carr=on :ace-#o-:ace in#erac#ions ?i#h each o: #he o#hers <eclines ?i#h #he num$er o: or-aniBa#ional )ar#ici)an#s. This cre a#es a shi:# in #he na#ure o: in#erac#ions such #ha# #he= assume a more im)ersonal, :ormal s#=le. 8lau an< a num$er o: coau#hors ha;e ar-ue< :or similar causal e::ec#s o: siBe on s#ruc#ure (8lau an< "co## 19 2, )). 222-!2I 8lau an< "choenherr 1971I 8lau 1972). If i# is #rue #ha# or-a niBa#ional :orm chan-es ?i#h siBe, selec#ion mechanisms ma= in<ee< o) era#e ?i#h re-ar< #o #he siBe <is#ri$u#ion. Fhen $i- or-aniBa#ions )re;ail i# ma= $e use:ul #o ;ie? #his as a s)ecial case o: selec#ion, in ?hich #he mo;emen# :rom Msmall :ormM #o Mlar-e :ormM is #heore#icall= in<is#in -uisha$le :rom #he <issolu#ion (M<ea#hM) o: small or-aniBa#ions an< #heir re)lacemen# $= (#he M$ir#hM o:) lar-e or-aniBa#ions. In sum, ?e ha;e i<en#i:ie< a num$er o: challen-es. The :irs# concerns #he #?o sources o: chan-e, selec#ion an< a<a)#i;e learnin-. Fe :eel #ha# #he or-aniBa#ions li#era#ure has o;erem)hasiBe< #he la##er a# #he eE)ense o: #he :ormer. Much more is +no?n a$ou# <ecision-ma+in- )rac#ices, :ore cas#in-, an< #he li+e #han a$ou# selec#ion in )o)ula#ions o: or-aniBa#ions. The secon< challen-e in;ol;es #he <is#inc#ion $e#?een selec#ion an< ;i a$ili#=. Fhe#her such a <is#inc#ion is necessar= <e)en<s on #he resul#s o: research on siBe ?hich is curren#l= $ein- )ursue< $= man= or-aniBa#ion researchers.
IV. TH6 5&IC4I5'6 7F I"7M7&5HI"M

In #he $es# <e;elo)e< s#a#emen# o: #he )rinci)les o: human ecolo-=, Ha?le= (19 8) ans?ers #he Gues#ion o: ?h= #here are so man= +in<s o: 170

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or-aniBa#ions. 3ccor<in- #o Ha?le=, #he <i;ersi#= o: or-aniBa#ional :orms is isomor)hic #o #he <i;ersi#= o: en;ironmen#s. In each <is#in-uisha$le en;ironmen#al con:i-ura#ion one :in<s, in eGuili$rium, onl= #ha# or-aniBa #ional :orm o)#imall= a<a)#e< #o #he <eman<s o: #he en;ironmen#. 6ach uni# eE)eriences cons#rain#s ?hich :orce i# #o resem$le o#her uni#s ?i#h #he same se# o: cons#rain#s. Ha?le=>s eE)lana#ion )laces hea;= em)hasis on communica#ion )a##erns an< s#ruc#ural com)lemen#s o: #hose )a##erns( MJ or-aniBa#ion uni#sK mus# su$mi# #o s#an<ar< #erms o: communica#ion an< #o s#an<ar< )roce<ures in conseGuence o: ?hich #he= <e;elo) similar in#ernal arran-emen#s ?i#hin limi#s im)ose< $= #heir res)ec#i;e siBesM (19 8, ). 22!). Fhile #he )ro)osi#ion seems com)le#el= soun< :rom an ecolo-ical )er s)ec#i;e, i# <oes no# a<<ress a num$er o: in#eres#in- consi<era#ions. There are a# leas# #?o res)ec#s in ?hich #he isomor)hism :ormula#ion mus# $e mo<i:ie< an< eE#en<e< i: i# is #o )ro;i<e sa#is:ac#or= ans?ers #o #he Gues #ion )ose<. The :irs# mo<i:ica#ion concerns #he mechanism or mechanisms res)onsi$le :or eGuili$rium. In #his res)ec#, #he )rinci)le o: isomor)hism mus# $e su))lemen#e< $= a cri#erion o: selec#ion an< a com)e#i#ion #heor=. The secon< mo<i:ica#ion <eals ?i#h #he :ac# #ha# #he )rinci)le o: iso mor)hism nei#her s)ea+s #o issues o: o)#imum a<a)#a#ion #o chan-inen;ironmen#s nor reco-niBes #ha# )o)ula#ions o: or-aniBa#ions o:#en :ace mul#i)le en;ironmen#s ?hich im)ose some?ha# inconsis#en# <eman<s. 3n un<ers#an<in- o: #he cons#rain#s on or-aniBa#ional :orms seems #o reGuire mo<elin- o: mul#i)le, <=namic en;ironmen#s. 7: course, ?e canno# :ull= eE#en< Ha?le=>s )rinci)le here. Fe a##em)# onl= #o ou#line #he main issues an< su--es# )ar#icular eE#ensions.
V. 47M56TITI7C TH67&9

The :irs# o: #he nee<e< eE#ensions is a s)eci:ica#ion o: #he o)#imiBa#ion )rocess res)onsi$le :or isomor)hism. Fe ha;e alrea<= <iscusse< #?o mech anisms( selec#ion an< a<a)#i;e learnin-. Isomor)hism can resul# ei#her $ecause nono)#imal :orms are selec#e< ou# o: a communi#= o: or-aniBa #ions or $ecause or-aniBa#ional <ecision ma+ers learn o)#imal res)onses an< a<,us# or-aniBa#ional $eha;ior accor<in-l=. Fe con#inue #o :ocus on #he :irs# o: #hese )rocesses( selec#ion. 4onsi<era#ion o: o)#imiBa#ion raises #?o issues( Fho is o)#imiBin-, an< ?ha# is $ein- o)#imiBe<. I# is Gui#e commonl= hel<, as in #he #heor= o: #he :irm, #ha# or-aniBa#ional <ecision ma+ers o)#imiBe )ro:i# o;er se#s o: or-aniBa#ional ac#ions. From a )o)ula#ion ecolo-= )ers)ec#i;e, i# is #he en;ironmen# ?hich o)#imiBes.M Fhe#her or no# in<i;i<ual or-aniBa5 In $iolo-ical a))lica#ions, one assumes #ha# )o?er (in #he )h=sical sense) is o)#imiBe< $= na#ural selec#ion in accor<ance ?i#h #he so-calle< Dar?in-'o#+a la?.

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#ions are consicousl= a<a)#in-, #he en;ironmen# selec#s ou# o)#imal com $ina#ions o: or-aniBa#ions. "o i: #here is a ra#ionali#= in;ol;e<, i# is #he Mra#ionali#=M o: na#ural selec#ion. 7r-aniBa#ional ra#ionali#= an< en;iron men#al ra#ionali#= ma= coinci<e in #he ins#ance o: :irms in com)e#i#i;e mar+e#s. In #his case, #he o)#imal $eha;ior o: each :irm is #o maEimiBe )ro:i# an< #he rule use< $= #he en;ironmen# (mar+e#, in #his case) is #o selec# ou# )ro:i# maEimiBers. Frie<man (1952) ma+es use o: #his o$ser

;a#ion #o )ro)ose a ,us#i:ica#ion o: #he #heor= o: #he :irm in #erms o: #he )rinci)les o: e;olu#ion. Ho?e;er, Fin#er (19 !) has ar-ue< con;incin-l= #ha# #he ac#ual si#ua#ion is much more com)lica#e< #han #his an< #ha# i# is mos# unusual :or in<i;i<ual ra#ionali#= an< en;ironmen#al or mar+e# ra#ionali#= #o lea< #o #he same o)#ima. Fhen #he #?o ra#ionali#ies <o no# a-ree, ?e are concerne< ?i#h #he o)#imiBin- $eha;ior o: #he en;ironmen#. 3 :ocus on selec#ion in;i#es an em)hasis on com)e#i#ion. 7r-aniBa#ional :orms )resuma$l= :ail #o :lourish in cer#ain en;ironmen#al circums#ances $ecause o#her :orms success:ull= com)e#e ?i#h #hem :or essen#ial resources. 3s lon- as #he resources ?hich sus#ain or-aniBa#ions are :ini#e an< )o)u la#ions ha;e unlimi#e< ca)aci#= #o eE)an<, com)e#i#ion mus# ensue. Ha?le= (1950, )). 201-2) :ollo?in- Dur+heim (19!7) amon- o#hers, )laces a hea;= em)hasis on com)e#i#ion as a <e#erminan# o: )a##erns o: social or-aniBa#ion. The <is#inc#i;e :ea#ure o: his mo<el is #he em)hasis on #he in<irec# na#ure o: #he )rocess( MThe ac#ion o: all on #he common su))l= -i;es rise #o a reci)rocal rela#ion $e#?een each uni# an< all #he o#hers, i: onl= :rom #he :ac# #ha# ?ha# one -e#s re<uces $= #ha# amoun# ?ha# #he o#hers can o$#ain ... ?i#hou# #his elemen# o: in<irec#ion, #ha# is, unless uni#s a::ec# one ano#her #hrou-h a::ec#in- a common limi#e< su))l=, com)e#i#ion <oes no# eEis#M (Ha?le= 1950, ). 202). In Ha?le=>s mo<el, com)e#i#ion )rocesses #=)icall= in;ol;e :our s#a-es( (1) <eman< :or resources eEcee<s su))l=I (2) com)e#i#ors $ecome more similar as s#an<ar< con<i#ions o: com)e#i#ion $rin- :or#h a uni:orm res)onseI (2) selec#ion elimina#es #he ?ea+es# com)e#i#orsI an< (!) <e)ose< com)e#i #ors <i::eren#ia#e ei#her #erri#oriall= or :unc#ionall=, =iel<in- a more com )leE <i;ision o: la$or. I# is sur)risin- #ha# #here is almos# no reliance on com)e#i#i;e mech anisms in Ha?le=>s la#er ?or+. In )ar#icular, as ?e no#e< a$o;e, #he ra#ionale -i;en :or #he isomor)hism )rinci)le uses an a<a)#a#ion lo-ic. Fe )ro)ose #o $alance #ha# #rea#men# $= a<<in- an eE)lici# :ocus on com)e#i#ion as a mechanism )ro<ucin- isomor)hism.M In so <oin-, ?e can $rin- a rich se# o: :ormal mo<els #o $ear on #he )ro$lem.
For #he case o: human social or-aniBa#ion, one mi-h# ar-ue #ha# selec#ion o)#imiBes #he u#iliBa#ion o: a s)eci:ic se# o: resources inclu<in- $u# no# res#ric#e< #o #he )o?er an< #he #ime o: mem$ers. Fe inclu<e onl= #he :irs# an< #hir< o: Ha?le=>" s#a-es in our mo<el o: com)e#i-

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The :irs# s#e) in cons#ruc#in- an ecolo-ical mo<el o: com)e#i#ion is #o s#a#e #he na#ure o: #he )o)ula#ion -ro?#h )rocess. 3# a minimum ?e ?ish #he mo<el #o incor)ora#e #he i<ea #ha# resources a;aila$le a# an= momen# :or each :orm o: or-aniBa#ion are :ini#e an< :iEe<. This corre s)on<s ?i#h Ha?le=>s no#ion o: limi#e< su))l= an< "#inchcom$e>s (19 5) ar-umen# #ha# human communi#ies ha;e limi#e< Mca)aci#ies :or or-aniB in-.M Fe also ?ish #o incor)ora#e #he ;ie? #ha# #he ra#e a# ?hich uni#s are a<<e< #o )o)ula#ions o: or-aniBa#ions <e)en<s on ho? much o: #he :iEe< ca)aci#= has alrea<= $een eEhaus#e<. The -rea#er #he uneEhaus#e< ca)aci#= in an en;ironmen#, #he :as#er shoul< $e #he ra#e o: -ro?#h o: )o)ula#ions o: or-aniBa#ions. 8u# #he ra#e a# ?hich )o)ula#ions o: or-a niBa#ions can eE)an< in#o unuse< ca)aci#= ;aries amon- :orms o: or-ani Ba#ion. "o #here are #?o <is#inc#i;e ecolo-ical consi<era#ions( #he ca)aci#= o: #he en;ironmen# #o su))or# :orms o: or-aniBa#ion an< #he ra#e a# ?hich #he )o)ula#ions -ro? (or <ecline) ?hen #he en;ironmen#al su))or# chan-es. In or<er #o s#a#e #he mo<el :ormall=, i# is hel):ul #o $e-in ?i#h #he con#rol :unc#ion #ha# Hummon, Doreian, an< Teu#er (1975) use #o a<< <=namic consi<era#ions #o 8lau>s #heor= o: siBe an< <i::eren#ia#ion. The con#rol mo<el s#a#es #ha# #he ra#e o: chan-e in #he siBe o: an= uni# (here a )o)ula#ion o: or-aniBa#ions) ;aries )ro)or#iona#el= ?i#h #he <i::erence $e#?een eEis#in- siBe, , an< #he eGuili$rium le;el o: siBe, !, )ermi##e<

in #ha# en;ironmen#. Then one )ossi$le re)resen#a#ion ?oul< $e

-== dt

(1)

"# ! $

==r# ! $

%.

In (1) ! an< r re)resen# #he limi#e< su))l= or en;ironmen#al ca)aci#= an< #he s#ruc#ural a$ili#= o: #he )o)ula#ion o: or-aniBa#ions #o res)on< #o chan-es in #he en;ironmen#, res)ec#i;el=. 3 )ar#icular :orm o: #he -eneral -ro?#h mo<el in (1) un<erlies mos# )o)ula#ion ecolo-= ?or+ on com)e#i#ion. This is #he lo-is#ic -ro?#h mo<el (:or )er ca)i#a -ro?#h)(

-d_

dt

==rl

s, k&

l%

(2)

?here l <eno#es )o)ula#ion siBe, k& is #he ca)aci#= o: #he en;ironmen# #o su))or# l (#his )arame#er is usuall= calle< #he carr=in- ca)aci#= ) , an< ri is #he so-calle< na#ural ra#e o: increase o: #he )o)ula#ion or #he ra#e a# ?hich #he )o)ula#ion -ro?s ?hen i# is :ar $elo? #he carr=in- ca)aci#=. 3s ?e in<ica#e< a$o;e, $o#h k an< r are ecolo-ical )arame#ers o: :un<a#ion. Fe )re:er #o #rea# uni:ormi#= o: res)onse an< communi#= <i;ersi#= as conse Guences o: com$ina#ions o: cer#ain com)e#i#i;e )rocesses an< en;ironmen#al :ea#ures.

172

men#al im)or#ance. 7ur research -rou) has $e-un #o com)are ;arious :orms o: or-aniBa#ion $= es#ima#in- #he )arame#ers o: mo<els li+e (2) :or each :orm o: or-aniBa#ion. Fe ha;e $een success:ul #o <a#e in rela#ins#ruc#ural :ea#ures o: or-aniBa#ions such as com)leEi#= o: core ac#i;i#= #o ;aria#ions in ran< k (Cielsen an< Hannan 1977I Freeman an< 8ri##ain

1977). This ?or+, #o-e#her ?i#h #ha# o: Hummon e# al. (1975), -i;es us con:i<ence #ha# #he mo<el in (1 ) an<*or (2) -i;es a -oo< a))roEima#ion o: #he -ro?#h o: )o)ula#ions o: or-aniBa#ions. %) #o #his )oin# ?e ha;e )resume< #ha# #he limi#s on -ro?#h re:lec# #he :ini#e na#ure o: #he en;ironmen# (e.-., communi#= ?eal#h an< miE o: occu)a#ional s+ills). I# is no? #ime #o rein#ro<uce com)e#i#ion. 3ccor<in#o Ha?le=, com)e#i#ion en#ers in<irec#l= ?hen #he com)e#i#ors lo?er #he :iEe< su))l=. Fe can mo<el #his $= :ollo?in- #he lea< o: $ioecolo-is#s an< eE#en<in- #he lo-is#ic -ro?#h mo<el. For eEam)le, consi<er a secon< )o)ula#ion o: or-aniBa#ions ?hose siBe is <eno#e< $= 2. The #?o )o)u la#ions are sai< #o com)e#e if #he a<<i#ion o: uni#s o: ei#her <ecreases #he ra#e o: -ro?#h o: #he o#her. This ?ill $e #he case ?hen $o#h )o)ula#ions are sus#aine< $= #he same #=)es o: resources. Then #he a))ro)ria#e mo<el is re)resen#e< in #he :ollo?in- s=s#em o: -ro?#h eGua#ions (+no?n as #he 'o#+a- Vol#erra eGua#ions :or com)e#in- )o)ula#ions) ( --==
dX 1

dt

r1P1

+l - Pl +l

lX12

X2)

(2 )

The coe::icien#s lX12 an< lX2b calle< com)e#i#ion coe::icien#s, <eno#e #he ma-ni#u<e o: #he e::ec# o: increases in one )o)ula#ion on #he -ro?#h o: #he o#her. In #his sim)le :ormula#ion, #he onl= conseGuence o: com)e#i#ion is #o lo?er #he carr=in- ca)aci#= o: #he en;ironmen# :or a )o)ula#ion o: or-aniBa#ions. 3nal=sis o: (2) )ro<uces in#eres#in- Guali#a#i;e resul#s. I# is no# <i::i cul# #o sho? #ha# a s#a$le #?o-)o)ula#ion eGuili$rium eEis#s :or #he s=s #em in (2) onl= i: (!) There:ore, ;er= similar )o)ula#ions (i.e., )o)ula#ions ?i#h com)e#i#ion coe::icien#s near uni#=) can coeEis# onl= un<er a ;er= )recise k'( +l ra#io. 3s a resul#, ?hen lX12 == lX21 == 1, no #?o-)o)ula#ion eGuili$rium can $e s#a$leI an= eEo-enous shoc+ ?ill resul# in #he elimina#ion o: one o: #he )o)ula#ions. This resul# su))or#s #he -enerali#= o: #he ?i<el= ci#e< M)rin-

ci)le o: com)e#i#i;e eEclusionM (Hause 192!).7 3ccor<in- #o #his )rinci)le, no #?o )o)ula#ions can con#inuousl= occu)= #he same niche. 5o)ula#ions are sai< #o occu)= #he same niche #o #he eE#en# #ha# #he= <e)en< on i<en#ical en;ironmen#al resources. If #he= are i<en#ical, #hen #he a<<i #ion o: an elemen# #o 2 has #he same conseGuences :or -ro?#h in I as <oes #he a<<i#ion o: an elemen# #o PlI in o#her ?or<s, #he com)e#i#ion coe::icien#s are uni#=. The $roa< conclusion is #ha# #he -rea#er #he simi lari#= o: #?o resource-limi#e< com)e#i#ors, #he less :easi$le is i# #ha# a sin-le en;ironmen# can su))or# $o#h o: #hem in eGuili$rium. If #?o )o)ula#ions o: or-aniBa#ions sus#aine< $= i<en#ical en;ironmen#al resources <i::er in some or-aniBa#ional charac#eris#ic, #ha# )o)ula#ion ?i#h #he charac#eris#ic less :i# #o en;ironmen#al con#in-encies ?ill #en< #o $e elimina#e<. The s#a$le eGuili$rium ?ill #hen con#ain onl= one )o)ula#ion ?hich can $e sai< #o $e isomor)hic #o #he en;ironmen#. In or<er #o see #he im)lica#ions o: #he mo<el :or or-aniBa#ional <i;ersi#=, ?e eE#en< #he 'o#+a-Vol#erra s=s#em #o inclu<e M com)e#i#ors(

-;It

dXi

// r.

, #ki

i $ )*ai+ +%(ki

(i

//

1, ... , ,%.

#-%

The -eneral s=s#em (5) has a communi#=

eGuili$rium( (i==l, ... ,M). ( )

These eGua#ions

can $e eE)resse< in ma#riE :orm( k

// A .,

(7)

?here . an< k are #, Pl) ma#riE(

column ;ec#ors an< A is #he communi#=

A== 1

amI

.QQ.

?hose elemen#s are #he com)e#i#ion coe::icien#s. The so-calle< #heor= o: communi#= s#ruc#ure en#ails #he anal=sis o: #he eGuili$rium $eha;ior o: #he s=s#em o: eGua#ion (7) :rom #he )ers)ec #i;e o: )os#ula#e< com)e#i#ion )rocesses.M The resul#s, #hou-h s#a#e< in
7 This so-calle< )rinci)le has mos#l= su--es#i;e ;alue (see Mac3r#hur J1972, )). !2! K :or a )ene#ra#in- cri#iGue o: a##em)#s #o <eri;e Guan#i#a#i;e im)lica#ions :rom Hause>s )rinci)leI mos# o: #hese cri#icisms <o no# a))l= #o #he Guali#a#i;e in:erences ?e consi<er). 8

Fe res#ric# a##en#ion #o #he case in ?hich all en#ries o: A are nonne-a#i;e. Ce-a-

3merican

Journal

o: "ociolo-=

#erms o: s)ecies <i;ersi#=, are Gui#e -eneral. In )ar#icular, one can sho? #ha# ?hen -ro?#h in )o)ula#ion is cons#raine< onl= $= resource a;ail a$ili#=, #he num$er o: <is#inc# resources se#s an u))er $oun< on <i;ersi#= in #he s=s#em.M 6;en more -enerall=, #he u))er $oun< on <i;ersi#= is eGual #o #he num$er o: <is#inc# resources )lus #he num$er o: a<<i#ional cons#rain#s on -ro?#h ('e;in 1970). It is <i::icul# #o a))l= ei#her resul# <irec#l= in or<er #o calcula#e #he u))er $oun< on <i;ersi#= e;en in #he nonhuman con#eE#. The chie: <i::i cul#= is #ha# o: i<en#i:=in- <is#inc# cons#rain#s. 3 -oo< <eal o: em)irical ?or+ is reGuire< i: one is #o ,u<-e ho? <i::eren# #?o cons#rain#s mus# $e in or<er #o ha;e <is#inc# conseGuences :or communi#= eGuili$ria. The #he orems <o, ho?e;er, im)l= use:ul Guali#a#i;e resul#s. If one can i<en#i:= en;ironmen#al chan-es ?hich a<< cons#rain#s #o a s=s#em or elimina#e #hem, one can conclu<e #ha# #he u))er $oun< o: <i;ersi#= is increase< or <ecrease<. This $roa< Guali#a#i;e resul# has a num$er o: )o#en#ial a))lica#ions #o #he research )ro$lems o: in#eres#. For eEam)le, #he eE)ansion o: mar +e#s an< o: s#a#e con#rol mechanisms #hrou-h social s=s#ems #en<s #o ha;e #he conseGuence o: elimina#in- or re<ucin- #he num$er o: cons#rain#s ?hich are i<ios=ncra#ic #o local en;ironmen#s. Vie?e< :rom #he )ers)ec #i;e o: #he lar-er s=s#em, #he )rocess o: eE)ansion o: #he economic an< )oli#ical cen#er shoul<, #hen, #en< #o re)lace some local cons#rain#s ?i#h more uni:orm ones. 3s lon- as #he local en;ironmen#s ?ere he#ero-eneous a# #he ou#se#, eE)ansion o: #he cen#er ou-h# #o re<uce #he num$er o: cons#rain#s on or-aniBa#ion in #he ?hole s=s#em. The #heor= ,us# <iscusse< im)lies on #he one han< #ha# #he chan-e in cons#rain# s#ruc#ure ou-h# #o lo?er or-aniBa#ional <i;ersi#= #hrou-h #he elimina#ion o: some )o)ula#ion.l. 7ne can ima-ine, on #he o#her han<, #ha# in some local en;ironmen#s, #he com$ina#ion o: unal#ere< local con s#rain#s an< ne? lar-er s=s#em cons#rain#s mi-h# increase #he #o#al num $er o: cons#rain#s in #he local s=s#em. In #ha# case, or-aniBa#ional <i;er si#= in #hose local en;ironmen#s shoul< increase. "uch an increase ?oul< resul# in #he crea#ion or a<o)#ion o: ne? or-aniBa#ional :orms. The increasin-l= im)or#an# role o: #he s#a#e in re-ula#in- economic an< social ac#ion )ro;i<es numerous o))or#uni#ies :or anal=Bin- #he im )ac# o: chan-es in cons#rain# s#ruc#ures on #he <i;ersi#= o: or-aniBa#ional
#i;e en#ries are a))ro)ria#e :or )re<a#or*)re= (or more -enerall=, hos#*)arasi#e) #ions. The #=)ical resul# :or #his case is c=clical )o)ula#ion -ro?#h. rela

9 3 more )recise s#a#emen# o: #he #heorem is #ha# no s#a$le eGuili$rium eEis#s :or a s=s#em o: M com)e#i#ors an< N M resources (Mac3r#hur an< 'e;ins 19 !). 10 For a more com)rehensi;e s#a#emen# o: #his ar-umen# ?i#h re:erence #o e#hnic or-aniBa#ion, see Hannan (1975).

<

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:orms. 4onsi<er #he im)ac# o: licensin- la?s, minimum ?a-e, heal#h, an< sa:e#= le-isla#ion, a::irma#i;e ac#ion, an< o#her re-ula#ions on or-aniBa #ional ac#ion. Fhen such re-ula#ions are a))lie< #o #he :ull ran-e o: or -aniBa#ions in $roa< areas o: ac#i;i#= #he= un<ou$#e<l= al#er #he siBe <is#ri$u#ions o: or-aniBa#ions. Mos# o:#en #he= selec# ou# #he smalles# or-aniBa#ions. 8u# i# is no# <i::icul# #o ima-ine si#ua#ions in ?hich me <ium-siBe< or-aniBa#ions (more )recisel=, #hose ?i#h some minimum le;el o: com)leEi#=) ?oul< $e more a<;ersel= a::ec#e<. 8esi<es al#erin- siBe <is#ri$u#ions, such re-ula#ions un<ou$#e<l= a::ec# #he <i;ersi#= o: or-a niBa#ional arran-emen#s in o#her ?a=s. Here one coul< anal=Be #he im)ac# o: s#a#e ac#ion on #he <i;ersi#= o: accoun#in- s=s#ems ?i#hin in<us#ries, curricula ?i#hin uni;ersi#ies, <e)ar#men#al s#ruc#ures ?i#hin hos)i#als, e#c. In each case i# ?oul< $e essen#ial #o <e#ermine ?he#her #he ne?l= im)ose< cons#rain# re)lace< lo?er le;el cons#rain#s, in ?hich case <i;er si#= shoul< <ecline, or ?he#her #he cons#rain# cumula#e< ?i#h #he eEis#incons#rain#s, in ?hich case or-aniBa#ional <i;ersi#= ?oul< $e li+el= #o increase. To in<ica#e #he richness o: #he sim)le com)e#i#ion #heor= ?e ha;e )ro)ose< ?e ?ill $rie:l= <iscuss ano#her sor# o: em)irical #es#. Fe no#e< a$o;e #ha# research on re-ula#ion mi-h# concern i#sel: ?i#h im)ac#s on <is#ri$u#ions o: or-aniBa#ions $= siBe. The classical mo<el o: or-aniBa #ional siBe <is#ri$u#ions ("imon an< 8onini 1958) )ro)oses #he :ollo?insim)le )rocess. 3 num$er o: or-aniBa#ions $e-in ?i#h #he same small siBe. "ome :rac#ion are a$le #o ma+e or $orro? some use:ul #echnical or or-a niBa#ional inno;a#ion #ha# )ermi#s #hem #o -ro? #o some lar-er siBe. Dur in- some s)eci:ie< #ime )erio< #he )rocess re)ea#s i#sel: ?i#h #he same :rac#ion ma+in- #he inno;a#ion reGuire< #o a##ain a lar-er siBe. "uch a -ro?#h )rocess e;en#uall= =iel<s #he lo-normal <is#ri$u#ion #ha# charac #eriBes so man= siBe <is#ri$u#ions. 4om)e#i#ion #heor= su--es#s a re:inemen# o: #his classical mo<el. If, as

?e ar-ue< earlier, lar-e chan-es in or-aniBa#ional siBe are accom)anie< $= s#ruc#ural chan-es (chan-es in :orm), or-aniBa#ions o: ;er= <i::eren# siBe in #he same area o: ac#i;i#= ?ill #en< #o eEhi$i# <i::eren# :orms. 3s a conseGuence o: #hese s#ruc#ural <i::erences, #he= ?ill #en< #o <e)en< on <i::eren# se#s o: en;ironmen#al resource (an< cons#rain#s). Tha# is, ?i#hin an= area o: ac#i;i#=, )a##erns o: resource use ?ill #en< #o $e s)ecialiBe< #o se-men#s o: #he siBe <is#ri$u#ion. This $ein- #he case, or-aniBa#ions ?ill com)e#e mos# in#ensel= ?i#h similar siBe or-aniBa#ions. 3lso, com)e#i#ion $e#?een )airs o: or-aniBa#ions ?i#hin an ac#i;i#= ?ill $e a <ecreasin:unc#ion o: #he <is#ance se)ara#in- #hem on #he siBe -ra<ien#. For eE am)le, small local $an+s com)e#e mos# ?i#h o#her small $an+s, #o a lesser eE#en# ?i#h me<ium-scale re-ional $an+s, an< har<l= a# all ?i#h in#erna#ional $an+s. %n<er #hese con<i#ions, si-ni:ican# al#era#ions in #he 177

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siBe <is#ri$u#ion in<ica#e selec#ion :or an< a-ains# cer#ain or-aniBa#ional :orms closel= associa#e< ?i#h re-ar< #o siBe. Co? le# us re#urn #o #he classical mo<el. Fhen lar-e-siBe< or-ani Ba#ions emer-e #he= )ose a com)e#i#i;e #hrea# #o me<ium-siBe< $u# har<l= an= #hrea# #o small or-aniBa#ions. In :ac#, #he rise o: lar-e or-aniBa#ions ma= increase #he sur;i;al chances o: small ones in a manner no# an#ici )a#e< in #he classical mo<el. Fhen #he lar-e or-aniBa#ions en#er, #hose in #he mi<<le o: #he siBe <is#ri$u#ion are #ra))e<. Fha#e;er s#ra#e-= #he= a<o)# #o :i-h# o:: #he challen-e o: #he lar-er :orm ma+es #hem more ;ulnera$le in com)e#i#ion ?i#h small or-aniBa#ions an< ;ice ;ersa. Tha# is, a# leas# in a s#a$le en;ironmen# #he #?o en<s o: #he siBe <is#ri$u#ion ou-h# #o ou#com)e#e #he mi<<le (see $elo?). "o in a lon-i#u<inal anal=sis o: or-aniBa#ional siBe <is#ri$u#ions ?e ?oul< eE)ec# #o see #he num$er o: me<ium-siBe< or-aniBa#ions <ecline u)on #he en#r= o: lar-er or-aniBa #ions. 3lso, ?e ?oul< eE)ec# #he :or#unes o: small or-aniBa#ions #o im )ro;e as #heir com)e#i#ors are remo;e< :rom #he en;ironmen#. This rea sonin- hol<s -enerall= :or com)e#i#ion alon- a sin-le -ra<ien#( #hose in #he mi<<le ?ill $e elimina#e< in s#a$le en;ironmen#s (Mac3r#hur 1972, )). !2-! ).
VI. CI4H6 TH67&9

The )rinci)le o: isomor)hism im)lies #ha# social or-aniBa#ions in eGui li$rium ?ill eEhi$i# s#ruc#ural :ea#ures #ha# are s)ecialiBe< #o salien# :ea #ures o: #he resource en;ironmen#. 3s lon- as #he en;ironmen# is s#a$le an< cer#ain, ?e see no <i::icul#= ?i#h #his )ro)osi#ion. 8u# <oes i# hol< ?hen #he en;ironmen# shi:#s ei#her )re<ic#a$l= or un)re<ic#a$l= amonse;eral al#erna#i;e con:i-ura#ions. Thou-h #he issues raise< $= a##em)# in- #o ans?er #his Gues#ion are com)leE, <oin- so is crucial #o <e;elo)ina<eGua#e mo<els o: or-aniBa#ional-en;ironmen# rela#ions. In#ui#ion su--es#s #ha# isomor)hism hol<s as a -oo< a))roEima#ion onl= in s#a$le en;ironmen#s. Face< ?i#h uns#a$le en;ironmen#s, or-ani Ba#ions ou-h# #o <e;elo) a -eneralis# s#ruc#ure #ha# is no# o)#imall= a<a)#e< #o an= sin-le en;ironmen#al con:i-ura#ion $u# is o)#imal o;er an en#ire se# o: con:i-ura#ions. In o#her ?or<s, ?e ou-h# #o :in< s)ecial iBe< or-aniBa#ions in s#a$le an< cer#ain en;ironmen#s an< -eneralis# or -aniBa#ions is uns#a$le an< uncer#ain en;ironmen#s. Fhe#her or no# #his sim)le )ro)osi#ion hol<s :or social or-aniBa#ions, onl= em)irical research ?ill #ell. Ho?e;er, a ;arie#= o: )o)ula#ion ecolo-= mo<els su--es#s #ha# i# is #oo sim)lis#ic. Fe canno# ho)e in one )a)er #o <e;elo) :ull= #he ar-umen#s in;ol;e<. Ins#ea< ?e in<ica#e #he main lines o: <e;elo)men# ?i#h re:erence #o one ra#her e;oca#i;e )ers)ec#i;e <e;elo)e< $= 'e;ins (19 2, 19 8)( #he #heor= o: niche ?i<#h. The conce)# o: Mniche,M ini#iall= $orro?e< $= $iolo-is#s :rom earl= 9!

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social science, )la=s a cen#ral role in ecolo-ical #heor=. This is no# #he )lace :or an eE#en<e< <iscussion o: #he mul#i)le uses o: #he conce)# (see Fhi##a+er an< 'e;in 197 ). The mo<el ?hich :ollo?s uses Hu#chinson>s ( 1957) :ormula#ion. From #his )oin# o: ;ie? #he (realiBe<) niche o: a )o)ula#ion is <e:ine< as #ha# area in cons#rain# s)ace (#he s)ace ?hose <imensions are le;els o: resources, e#c.) in ?hich #he )o)ula#ion ou#com )e#es all o#her local )o)ula#ions. The niche, #hen, consis#s o: all #hose com$ina#ions o: resource le;els a# ?hich #he )o)ula#ion can sur;i;e an< re)ro<uce i#sel:. 6ach )o)ula#ion occu)ies a <is#inc# niche. For )resen# )ur)oses i# su: :ices #o consi<er cases ?here )airs o: )o)ula#ions <i::er ?i#h res)ec# #o a sin-le en;ironmen#al <imension, E, an< are ali+e ?i#h res)ec# #o all

o#hers. Then rela#i;e com)e#i#i;e )osi#ions can $e sim)l= summariBe< as in :i-ure 1. 3s ?e ha;e <ra?n #his :i-ure, one )o)ula#ion, A, occu)ies a

Fitness (F)

FIH. 1.-Fi#ness :unc#ions (niches) :or s)ecialis#s an< -eneralis#s

;er= $roa< niche, ?hereas #he o#her, B, has concen#ra#e< i#s :i#ness, <e no#e< /, on a ;er= narro? $an< o: en;ironmen#al ;aria#ion. This <is #inc#ion, ?hich is usuall= re:erre< #o as -eneralism ;ersus s)ecialism, is crucial #o $iolo-ical ecolo-= an< #o a )o)ula#ion ecolo-= o: or-aniBa#ions. 179

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In essence, #he <is#inc#ion $e#?een s)ecialism an< -eneralism re:ers #o ?he#her a )o)ula#ion o: or-aniBa#ions :lourishes $ecause i# maEimiBes i#s eE)loi#a#ion o: #he en;ironmen# an< acce)#s #he ris+ o: ha;in- #ha# en ;ironmen# chan-e or $ecause i# acce)#s a lo?er le;el o: eE)loi#a#ion in re#urn :or -rea#er securi#=. Fhe#her or no# #he eGuili$rium <is#ri$u#ion o: or-aniBa#ional :orms is <omina#e< $= #he s)ecialis# <e)en<s, as ?e ?ill see, on #he sha)e o: #he :i#ness se#s an< on )ro)er#ies o: #he en;ironmen#. 5ar# o: #he e::icienc= resul#in- :rom s)ecialism is <eri;e< :rom #he lo?er reGuiremen#s :or eEcess ca)aci#=. Hi;en some uncer#ain#=, mos# or-aniBa#ions main#ain some eEcess ca)aci#= #o insure #he relia$ili#= o: )er:ormance. In a ra)i<l= chan-in- en;ironmen#, #he <e:ini#ion o: eEcess ca)aci#= is li+el= #o chan-e :reGuen#l=. Fha# is use< #o<a= ma= $ecome eEcess #omorro?, an< ?ha# is eEcess #o<a= ma= $e crucial #omorro?. 7r-aniBa#ions o)era#in- in en;ironmen#s ?here #he #ransi#ion :rom s#a#e #o s#a#e is less :reGuen# ?ill (in eGuili$rium) ha;e #o main#ain eEcess ca)aci#= in a -i;en alloca#ional )a##ern :or lon-er )erio<s o: #ime. Fhereas #hose char-e< ?i#h assessin- )er:ormance ?ill $e #em)#e< #o ;ie? such alloca#ions as ?as#e:ul, #he= ma= $e essen#ial :or sur;i;al. Thom)son (19 7) has ar-ue< #ha# or-aniBa#ions alloca#e resources #o uni#s char-e< ?i#h #he :unc#ion o: insula#in- core #echnolo-= :rom en;ironmen#all= in <uce< <isru)#ion. "o, :or eEam)le, manu:ac#urin- :irms ma= re#ain or em)lo= le-al s#a::s e;en ?hen #he= are no# curren#l= :acin- li#i-a#ion. The im)or#ance o: eEcess ca)aci#= is no# com)le#el= $oun< u) ?i#h #he issue o: ho? much eEcess ca)aci#= ?ill $e main#aine<. I# also in;ol;es #he manner in ?hich i# is use<. 7r-aniBa#ions ma= insure relia$le )er :ormance $= crea#in- s)ecialiBe< uni#s, as Thom)son (19 7) su--es#s, or #he= ma= alloca#e eEcess ca)aci#= #o or-aniBa#ional roles, $= em)lo=in)ersonnel ?i#h s+ills an< a$ili#ies ?hich eEcee< #he rou#ine reGuiremen#s o: #heir ,o$s. This is one o: #he im)or#an# reasons :or usin- )ro:essionals in or-aniBa#ions. 5ro:essionals use mRJre resources no# onl= $ecause #he= #en< #o $e )ai< more, $u# also $ecause or-aniBa#ions mus# allo? #hem more <iscre#ion (inclu<in- #he :ree<om #o res)on< #o ou#si<e re:erence -rou)s). 7r-aniBa#ions, in #urn, $ecome more :leEi$le $= em)lo=in- )ro :essionals. The= increase #heir ca)aci#= #o <eal ?i#h a ;aria$le en;iron men# an< #he con#in-encies i# )ro<uces. For eEam)le, hos)i#als an< #heir )a#ien#s o:#en em)lo= o$s#e#ricians an< )e<ia#ricians in #heir <eli;er= rooms e;en #hou-h #he normal <eli;er= o: $a$ies can $e )er:orme< eGuall= ?ell, an< )erha)s e;en $e##er, $= mi<?i;es. The s+ills o: #he me<ical <oc#or re)resen# eEcess ca)aci#= #o insure relia$le )er:ormance shoul< <eli;er= no# $e normal. %suall=, #he )e<ia#rician eEamines #he in:an# imme<ia#el= a:#er $ir#h #o see i: #here is an= a$normali#= reGuirin- imme <ia#e ac#ion. If #he mo#her is su::erin- <an-erous conseGuences :rom -i;

in- $ir#h, an< #he chil< is also in nee< o: a##en#ion, #he )resence o: #he 180

5o)ula#ion )e<ia#rician insures #ha# #he o$s#e#rician #hem in alloca#in- his a##en#ion.

6colo-= o: 7r-aniBa#ions

?ill no# ha;e #o choose $e#?een

6Ecess ca)aci#= ma= also $e alloca#e< #o #he <e;elo)men# an< main #enance o: )roce<ural s=s#ems. Fhen #he cer#ain#= o: a -i;en en;iron men#al s#a#e is hi-h, or-aniBa#ional o)era#ions shoul< $e rou#ine, an< co or<ina#ion can $e accom)lishe< $= :ormaliBe< rules an< #he in;es#men# o: resources in #rainin- incum$en#s #o :ollo? #hose :ormaliBe< )roce<ures. If in :ac# #he en;ironmen# ?ere unchan-in- (p 1), all )ar#ici)an#s

//

?ere )roce<urall= s+ille<, an< #he )roce<ures ?ere )er:ec#l= #une<, #here ?oul< $e no nee< :or an= con#rol s#ruc#ure a# all, eEce)# #o moni#or $eha;ior. Ho?e;er, ?hen cer#ain#= is lo?, or-aniBa#ional o)era#ions are less rou#ine. %n<er #hese circums#ances, a -rea#er alloca#ion o: resources #o <e;elo) an< main#ain )roce<ural s=s#ems is coun#er)ro<uc#i;e an< o)#imal or-aniBa#ional :orms ?ill alloca#e resources #o less :ormaliBe< s=s#ems ca)a$le o: more inno;a#i;e res)onses (e.-., commi##ees an< #eams). In #his case, eEcess ca)aci#= is re)resen#e< $= #he increase< #ime i# #a+es such s#ruc#ures #o ma+e <ecisions an< $= increase< coor<ina#ion cos#s. The )oin# here is #ha# )o)ula#ions o: or-aniBa#ional :orms ?ill $e selec#e< :or or a-ains# <e)en<in- u)on #he amoun# o: eEcess ca)aci#= #he= main#ain an< ho? #he= alloca#e i#. I# ma=or ma= no# $e ra#ional :or an= )ar#icular or-aniBa#ion #o a<o)# one )a##ern or ano#her. Fha# ?oul< seem li+e ?as#e #o an=one assessin- )er:ormance a# one #ime ma= $e #he <i::erence $e#?een sur;i;al an< :ailure la#er. "imilarl=, or-aniBa #ions ma= sur;i;e $ecause hi-h le;els o: )ro:essionaliBa#ion )ro<uce co or<ina#ion $= mu#ual a<,us#men# <es)i#e a some?ha# chao#ic a))earance. 7#hers, in ?hich e;er=one seems #o +no? )recisel= ?ha# he is <oin- a# all #imes, ma= :ail. %n<er a -i;en se# o: en;ironmen#al circums#ances #he :un<amen#al ecolo-ical Gues#ion is( ?hich :orms #hri;e an< ?hich :orms <isa))ear. Heneralism ma= $e o$ser;e< in a )o)ula#ion o: or-aniBa#ions, #hen, ei#her in i#s reliance u)on a ?i<e ;arie#= o: resources simul#aneousl= or in i#s main#enance o: eEcess ca)aci#= a# an= -i;en #ime. This eEcess ca )aci#= allo?s such or-aniBa#ions #o chan-e in or<er #o #a+e a<;an#a-e o: resources ?hich $ecome more rea<il= a;aila$le. 4or)ora#ions ?hich main #ain an unusuall= lar-e )ro)or#ion o: #heir #o#al asse#s in :lui< :orm (Mslac+,M in #erms o: #heor= o: #he :irmI 5enrose 1959I 4=er# an< March 19 2) are -eneraliBin-. In ei#her case, -eneralism is cos#l=. %n<er s#a$le en;ironmen#al circums#ances, -eneralis#s ?ill $e ou#com)e#e< $= s)ecial is#s. 3n< a# an= -i;en )oin# in #ime, a s#a#ic anal=sis ?ill re;eal eEcess ca)aci#=. 3n im)lica#ion-shi:#in- our :ocus #o in<i;i<ual -eneralis#s-is #ha# ou#si<e a-en#s ?ill o:#en mis#a+e eEcess ca)aci#= :or ?as#e. 181

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Fe can in;es#i-a#e #he e;olu#ion o: niche ?i<#h i: ?e ma+e #he as sum)#ion #ha# areas un<er #he :i#ness cur;e are eGual, an< #ha# s)ecialis#s <i::er :rom -eneralis#s in ho? #he= <is#ri$u#e #he :iEe< Guan#i#= o: :i#ness o;er en;ironmen#al ou#comes. ")ecialis#s ou#com)e#e -eneralis#s o;er #he ran-e o: ou#comes #o ?hich #he= ha;e s)ecialiBe< ($ecause o: #he :iEe< le;el o: :i#ness assum)#ion). 3s lon- as #he en;ironmen#al ;aria#ion re mains ?i#hin #ha# in#er;al (#he in#er;al [m,n] in :i-. 1), -eneralis#s ha;e no a<a)#i;e a<;an#a-e an< ?ill $e selec#e< a-ains#. 3l#erna#i;el=, if #he

en;ironmen# is onl= occasionall= ?i#hin #he in#er;al, s)ecialis#s ?ill :are less ?ell #han -eneralis#s. These $rie: commen#s ma+e clear #he im)or #ance o: en;ironmen#al ;aria#ion :or #he e;olu#ion o: niche ?i<#h. To sim)li:= :ur#her, consi<er an en;ironmen# ?hich can #a+e on onl= #?o s#a#es an< in e;er= )erio< :alls in s#a#e one ?i#h )ro$a$ili#= p an< in s#a#e #?o ?i#h )ro$a$ili#= q // (1 - p). 3ssume :ur#her #ha# ;aria #ions in en;ironmen#al s#a#es are 8ernoulli #rials (in<e)en<en# :rom )e rio< #o )erio<). For #his si#ua#ion 'e;ins (19 2, 19 8) has sho?n #ha# o)#imal niche ?i<#h <e)en<s on p an< #he M<is#anceM $e#?een #he #?o s#a#es o: #he en;ironmen#. To see #his, ?e chan-e :ocus sli-h#l=. "ince each or-aniBa#ion :aces #?o en;ironmen#s, i#s :i#ness <e)en<s on :i#ness in #he )air. Fe can sum mariBe #he a<a)#i;e )o#en#ial o: each or-aniBa#ion $= )lo##in- #hese )airs o: ;alues (:i#ness in s#a#e 1 an< in s#a#e 2) in a ne? s)ace ?hose aEes are :i#ness in each o: #he s#a#es, as in :i-ure 2. In #his re)resen#a#ion, each )oin# <eno#es #he :i#ness o: a <is#inc# or-aniBa#ional :orm. The clou< o: )oin#s is #erme< #he M:i#ness se#.M Fe )resume #ha# all o: #he na#urall= )ossi$le a<a)#a#ions are re)resen#e< in #he :i#ness se#. 7ur in#eres# is in <e#erminin- ?hich )oin#s in #he :i#ness se# ?ill $e :a;ore< $= na#ural selec#ion. Co#ice :irs# #ha# all )oin#s in#erior #o #he se# are in:erior in #erms o: :i#ness #o a# leas# some )oin# on #he $oun<ar= o: #he se#. In #his sense #he $oun<ar=, <ra?n as a con#inuous line, re) resen#s #he o)#imal )ossi$ili#ies. "ince na#ural selec#ion maEimiBes :i#ness, i# mus# choose )oin#s on #he $oun<ar=. This narro?s our search #o see+ in- ?hich :orm (s) on #he $oun<ar= ?ill $e :a;ore<. 3s :i-ure 'b is <ra?n, no or-aniBa#ional :orm <oes )ar#icularl= ?ell in $o#h s#a#es o: #he en;ironmen#-no :orm has hi-h le;els o: :i#ness in $o#h. This ?ill $e #he case ?hen #he #?o s#a#es are M:ar a)ar#M in #he sense #ha# #he= im)ose ;er= <i::eren# a<a)#i;e con#in-encies on or-ani Ba#ions. In such cases (see 'e;ins 19 8), #he :i#ness se# ?ill $e conca;e. Fhen #he M<is#anceM $e#?een s#a#es is small, #here is no reason ?h= cer #ain or-aniBa#ional :orms canno# <o ?ell in $o#h en;ironmen#s. In such cases, #he :iness se# ?ill $e con;eE, as in :i-ure 'a. The :i#ness :unc#ions in :i-ures 'a an< 'b <escri$e <i::eren# a<a)#i;e si#ua#ions. The neE# s#e) is #o mo<el #he o)#imiBa#ion )rocess. To <o so, 182

5o)ula#ion

6colo-= o: 7r-aniBa#ions

-,

-, M )/l.7

.... .. '. ~-,---------

R>))I>SiI(1(RI~(( ;.>' "

<: :........
... :::.' :..
"

p=, 5

-,

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-,
>T. )/.5 -, <,

b FI . 2.-0)#imal a<a)#a#ion in :ine--raine< en;ironmen#I a, con;eE :i#ness se#I b, conca;e :i#ness se#.

?e in#ro<uce a :ur#her <is#inc#ion. 6colo-is#s ha;e :oun< i# use:ul #o <is#in-uish $o#h s)a#ial an< #em)oral en;ironmen#al ;aria#ion accor<in#o -rain. 6n;ironmen#al ;aria#ion is sai< #o $e :ine--raine< ?hen a #=) ical elemen# (or-aniBa#ion) encoun#ers man= uni#s or re)lica#ions. From 182

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a #em)oral )ers)ec#i;e, ;aria#ion is :ine--raine< ?hen #=)ical <ura#ions in s#a#es are shor# rela#i;e #o #he li:e#ime o: or-aniBa#ions. 7#her?ise, #he en;ironmen# is sai< #o $e coarse--raine<. Deman< :or )ro<uc#s or ser ;ices is o:#en charac#eriBe< $= :ine--raine< ;aria#ion ?hereas chan-es in le-al s#ruc#ures are more #=)icall= coarse--raine<. The essen#ial <i::erence $e#?een #he #?o #=)es o: en;ironmen#al ;aria #ion is #he cos# o: su$o)#imal s#ra#e-ies. The )ro$lem o: ecolo-ical a<a) #a#ion can $e consi<ere< a -ame o: chance in ?hich #he )o)ula#ion chooses a s#ra#e-= (s)ecialism or -eneralism) an< #hen #he en;ironmen# chooses an ou#come ($=, sa=, :li))in- a coin). If #he en;ironmen# Mcomes

u)M in a s#a#e :a;ora$le #o #he or-aniBa#ional :orm, i# )ros)ersI o#her?ise, i# <eclines. Ho?e;er, i: #he ;aria#ion is :ine--raine< (<ura#ions are shor#), each )o)ula#ion o: or-aniBa#ions eE)eriences a -rea# man= #rials an< en ;ironmen# is eE)erience< as an a;era-e. Fhen ;aria#ion is coarse--raine<, ho?e;er, #he )erio< o: <ecline s#emmin- :rom a ?ron- choice ma= eEcee< #he or-aniBa#ional ca)aci#= #o sus#ain i#sel: un<er un:a;ora$le con<i#ions. To ca)#ure #hese <i::erences, 'e;ins in#ro<uce< an a<a)#i;e :unc#ion #o re)resen# ho? na#ural selec#ion ?oul< ?ei-h# :i#ness in each s#a#e un<er #he <i::eren# con<i#ions. In <iscussin- :ine--raine< ;aria#ion, ?e su--es#e< #ha# #he en;ironmen# is eE)erience< as an a;era-e.lO The a) )ro)ria#e a<a)#i;e :unc#ion, #hen, sim)l= ?ei-h#s :i#ness in #he #?o s#a#es (FI an< W'% accor<in- #o :reGuenc= o: occurrence( A (W*,W'% == pW* qW'0 In or<er #o consi<er o)#imal a<a)#a#ion ?e merel=

su )erim)ose #he a<a)#i;e :unc#ion on #he :i#ness se# an< :in< )oin#s o: #an-enc= o: a<a)#i;e :unc#ion an< :i#ness :unc#ions. 5oin#s o: #an-enc= are o)#imal a<a)#a#ions. The solu#ions :or ;arious cases are )resen#e< in :i-ure 2. If #he en;ironmen# is com)le#el= s#a$le (i.e., p == 1), #hen s)e cialism is o)#imal. If #he en;ironmen# is maEimall= uncer#ain (i.e., 1 == .5), -eneralism is o)#imal in #he con;eE case (?hen #he <eman<s o: #he <i::eren# en;ironmen#s are no# #oo <issimilar) $u# no# in #he conca;e case. In :ac#, as #he mo<el is <e;elo)e<, s)ecialism al?a=s ?ins ou# in #he conca;e case. 4onsi<er :irs# #he cases in ?hich #he en;ironmen# is s#a$le ('e., p == 1). Co# sur)risin-l=, s)ecialism is o)#imal. The resul#s :or uns#a$le en ;ironmen#s <i;er-e. Fhen #he :i#ness se# is con;eE (i.e., #he <eman<s o: #he <i::eren# en;ironmen#al s#a#es are similar an<*or com)lemen#ar=), -eneralism is o)#imal. 8u# ?hen #he en;ironmen#al <eman<s <i::er (an< #he :i#ness se# is conca;e), s)ecialism is o)#imal. This is no# as s#ran-e
11 Tha# selec#ion <e)en<s on a;era-e ou#comes is onl= one h=)o#hesis. Tem)le#on an< &o#hman (197!) ar-ue #ha# selec#ion <e)en<s no# on a;era-e ou#comes $u# on some minimum le;el o: :i#ness. Fhe#her a;era-e ou#comes or some o#her cri#erion -ui<es selec#ion in )o)ula#ions o: or-aniBa#ions is o)en #o Gues#ion. Fe :ollo? 'e;ins

18!

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in or<er #o +ee) #he eE)osi#ion sim)le.

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a resul# as i# :irs# a))ears. Fhen #he en;ironmen# chan-es ra)i<l= amonGui#e <i::eren# s#a#es, #he cos# o: -eneralism is hi-h. "ince #he <eman<s in #he <i::eren# s#a#es are <issimilar, consi<era$le s#ruc#ural mana-emen# is reGuire< o: -eneralis#s. 8u# since #he en;ironmen# chan-es ra)i<l=, #hese or-aniBa#ions ?ill s)en< mos# o: #heir #ime an< ener-ies a<,us#ins#ruc#ure. I# is a))aren#l= $e##er un<er such con<i#ions #o a<o)# a s)ecial iBe< s#ruc#ure an< Mri<e ou#M #he a<;erse en;ironmen#s. The case o: coarse--raine< en;ironmen#s is some?ha# more com)leE. 7ur in#ui#i;e un<ers#an<inis #ha# since #he <ura#ion o: an en;iron men#al s#a#e is lon-, mala<a)#a#ion ou-h# #o $e -i;en -rea#er ?ei-h#. Tha# is, #he cos#s o: mala<a)#a#ion -rea#l= ou#?ei-h an= a<;an#a-e in

me#ho< o: :in<in- o)#imal a<a)#a#ions is #he same. The resul#s are :oun< in :i-ure 2. 7nl= one case <i::ers :rom ?ha# ?e :oun< :or :ine--raine< en;ironmen#s( #he com$ina#ion o: uncer#ain an< coarse--raine< ;aria#ion ?i#h conca;e :i#ness se#s. Fe sa? a$o;e #ha# ?hen such ;aria#ion is :ine -raine<, i# is $e##er #o s)ecialiBe. Fhen #he <ura#ion o: en;ironmen#al s#a#es is lon-, ho?e;er, #he cos#s o: #his s#ra#e-= are -rea#. 'on- )erio<s o: nona<a)#a#ion ?ill #hrea#en #he sur;i;al o: #he or-aniBa#ion. In a<<i #ion, #he :ac# #ha# #he en;ironmen# chan-es less o:#en means #ha# -en eralis#s nee< no# s)en< mos# o: #heir #ime an< ener-ies al#erin- s#ruc#ure. Thus -eneralism is #he o)#imal s#ra#e-= in #his case as ?e see in :i-ure 2b. The com$ina#ion o: coarse--raine< en;ironmen#al ;aria#ion an< con ca;e :i#ness se#s raises a :ur#her )ossi$ili#=. The o)#imal a<a)#a#ion in #he :ace o: en;ironmen#al uncer#ain#= )ossesses :airl= lo? le;els o: :i#ness in ei#her s#a#e. I# seems clear #ha# #here mus# $e a $e##er solu#ion. 'e;ins <iscusses #his case in <e)#h an< conclu<es #ha# :or #he $iolo-ical case ?i#h -ene#ic #ransmission o: s#ruc#ure M)ol=mor)hismM or -ene#icall= main#aine< )o)ula#ion he#ero-enei#= ?ill $e selec#e< :or. The su--es#ion is #ha# )o)ula#ions com$ine #=)es (<i::erin-, sa=, in color, $loo< #=)e, e#c.) some o: ?hich are s)ecialiBe< #o s#a#e 1 an< some #o s#a#e 2. Fi#h such a com$ina#ion a# leas# a )or#ion o: #he )o)ula#ion ?ill al?a=s :lour ish an< main#ain #he -ene#ic <i;ersi#= ?hich allo?s i# #o con#inue #o :lourish ?hen #he en;ironmen# chan-es s#a#e. The se# o: all such he#ero -eneous )o)ula#ions (com)ose< o: )ro)or#ions o: s)ecialis#s #o each o: #he #?o en;ironmen#s) can $e re)resen#e< in #he :i#ness <ia-rams as a s#rai-h# line ,oinin- #he mos# eE#reme )oin#s ?i#h all com$ina#ions :allin?i#hin #his line. 4oarse--raine< an< uncer#ain ;aria#ion :a;ors a <is#inc# :orm o: -en eralism( )ol=mor)hism. Fe <o no# ha;e #o search ;er= :ar #o :in< an analo-ous ou#come. 7r-aniBa#ions ma= :e<era#e in such a ?a= #ha# su)ra or-aniBa#ions consis#in- o: he#ero-eneous collec#ions o: s)ecialis# or-ani-

curre< $= #he correc# choice. 7ne a<a)#i;e :unc#ion ?hich -i;es #his resul# is #he lo--linear mo<el use< $= 'e;ins( A (Wl,W'% W& W'!. The

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Ba#ions )ool resources. Fhen #he en;ironmen# is uncer#ain an< coarse -raine< an< su$uni#s <i::icul# #o se# u) an< #ear <o?n, #he cos#s o: main #ainin- #he un?iel<= s#ruc#ure im)ose< $= :e<era#ion ma= $e more #han o::se# $= #he :ac# #ha# a# leas# a )or#ion o: #he amal-ama#e< or-aniBa#ion ?ill <o ?ell no ma##er ?ha# #he s#a#e o: #he en;ironmen#. In #erms o: #he mo<el su--es#e< a$o;e #here are no o#her si#ua#ions in ?hich such 187

:e<era#e< or-aniBa#ions ha;e a com)e#i#i;e a<;an#a-e. 3n< e;en in #his case, #he onl= #ime <urin- ?hich #he= ha;e such an a<;an#a-e is ?hen coarse--raine< ;aria#ion is uncer#ain. "uch an amal-ama#e< Mhol<in- com)an=M )a##ern ma= $e o$ser;e< in mo<ern uni;ersi#ies. 6nrollmen# an< research su))or# ?aE an< ?ane o;er #ime as <o #he =iel< on in;es#e< en<o?men# securi#ies an< #he $ene:icence o: le-isla#ures. "ome o: #hese resources :ollo? )re<ic#a$le c=cles. 7#hers <o no#. 8u# it is eE#remel= eE)ensi;e #o $uil< u) an< <isman#le aca<emic uni#s. It is cos#l= no# onl= in mone= $u# also in #he ener-ies consume<

$= )oli#ical con:lic#. 4onseGuen#l=, uni;ersi#ies are cons#an#l= M#aEin-M su$uni#s ?i#h )len#i:ul en;ironmen#s #o su$si<iBe less :or#una#e su$uni#s. It is common, :or ins#ance, :or uni;ersi#ies #o alloca#e :acul#= )osi#ions accor<in- #o some :iEe< mas#er )lan, un<ersu))or#in- #he ra)i<l= -ro?in<e)ar#men#s an< main#ainin- eEcess :acul#= in o#hers. This )ar#ial eE)la na#ion o: #he un?iel<l= s#ruc#ures #ha# encom)ass li$eral ar#s <e)ar#men#s, )ro:essional schools, research la$ora#ories, e#c., is a# leas# as )ersuasi;e as eE)lana#ions #ha# em)hasiBe in#ellec#ual in#er<e)en<ence amon- uni#s. Much more can $e sai< concernin- a))lica#ions o: niche #heor= #o or-aniBa#ion-en;ironmen# rela#ions. Fe ha;e :ocuse< on a sim)le ;ersion hi-hli-h#in- #he in#er)la= $e#?een com)e#i#ion an< en;ironmen#al ;aria #ion in #he <e#ermina#ion o: o)#imal a<a)#i;e s#ruc#ure in or<er #o sho? #ha# #he )rinci)le o: isomor)hism nee<s consi<era$le eE)ansion #o <eal ?i#h mul#i)le en;ironmen#al ou#comes an< #heir associa#e< uncer#ain#=. The li#era#ure in ecolo-= #o ?hich ?e ha;e ma<e re:erence is -ro?ineE)onen#iall= a# #he momen# an< ne? resul#s an< mo<els are a))earinmon#hl=. The )ro<uc#s o: #hese <e;elo)men#s )ro;i<e s#u<en#s o: or-ani Ba#ions ?i#h a rich )o#en#ial :or #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions. 4onsi<er an eEam)le. In his anal=sis o: $ureaucra#ic an< cra:# a<min is#ra#ion or )ro<uc#ion, "#inchcom$e (1959) ar-ue< #ha# cons#ruc#ion :irms <o no# rel= u)on $ureaucra#icall= or-aniBe< a<minis#ra#i;e s#a::s $ecause o: seasonal :luc#ua#ions in <eman<. 3<minis#ra#i;e s#a::s cons#i #u#e an o;erhea< cos# ?hich remains rou-hl= cons#an# o;er #he =ear. The a<;an#a-e o: #he o#her?ise cos#l= (in #erms o: salaries) cra:# a<minis#ra #ion is #ha# coor<ina#ion o: ?or+ is accom)lishe< #hrou-h a reliance u)on )rior socialiBa#ion o: cra:#smen an< u)on or-aniBa#ion. "ince em)lo=men# le;els can more easil= $e increase< or <ecrease< ?i#h <eman< un<er a cra:# s=s#em, a<minis#ra#i;e cos#s are more easil= al#ere< #o mee# <eman<. The :un<amen#al source o: #his )a##ern is #he seasonal ;aria#ion in cons#ruc#ion. In ecolo-ical #erms, #he <eman< en;ironmen# is coarse -raine<. In a<<i#ion, #he #?o s#a#es <e:ine< $= season are Gui#e <i::eren#, resul#in- in a conca;e :i#ness cur;e. 4ra:#-a<minis#ere< housin- cons#ruc #ion :irms are )ro$a$l= Gui#e ine::icien# ?hen <eman< is a# i#s )ea+ an<

?hen #he +in< o: housin- un<er cons#ruc#ion is s#an<ar<iBe<. In such si#ua#ions, ?e ?oul< eE)ec# #his :orm o: or-aniBa#ion #o :ace s#i:: com )e#i#ion :rom o#her :irms. For ins#ance, in re-ions ?here housin- con s#ruc#ion is less seasonal, mo<ular housin-, mo$ile homes, an< )re:a$ri ca#e< housin- are more li+el= #o :lourish an< ?e ?oul< eE)ec# #he con s#ruc#ion $usiness #o $e more hi-hl= $ureaucra#iBe<. 3no#her ;aria#ion in <eman< is #o $e :oun< in #he $usiness c=cle. Fhile seasonal :luc#ua#ions are s#a$le (uncer#ain#= is lo?), in#eres# ra#es, la$or rela#ions, an< ma#erials cos#s are more <i::icul# #o )re<ic#. Varia#ions o: #his sor# shoul< :a;or a -eneralis# mo<e o: a<a)#a#ion. Tha# is, ?hen en ;ironmen#s are coarse--raine<, charac#eriBe< $= conca;e :i#ness cur;es, an< uncer#ain, )o)ula#ions o: or-aniBa#ions ?ill $e more li+el= #o sur;i;e i: #he= he<-e #heir $e#s $= see+in- a ?i<er ;arie#= o: resource $ases. For #his reason, ?e #hin+, cra:#-a<minis#ere< cons#ruc#ion or-aniBa#ions are :reGuen#l= -eneral con#rac#ors ?ho no# onl= $uil< houses $u# en-a-e in o#her +in<s o: cons#ruc#ion as ?ell (sho))in- )laBas, o::ice $uil<in-s, e#c.). In com)arison, mo<ular housin- is chea)er an< #he uni#s are in s#alle< on ren#e< s)ace. 4onseGuen#l=, in#eres# ra#es are less im)or#an#. "ince or-aniBa#ions )ro<ucin- #his +in< o: housin- <o no# em)lo= cra:#s men $u# use #he chea)es# an< leas# s+ille< la$or #he= can o$#ain, la$or rela#ions are less )ro$lema#ical. I# ma= also $e #ha# #heir reliance on <i::eren# ma#erials (e.-., shee# aluminum) con#ri$u#es #o a lo?er le;el o: uncer#ain#=. In conseGuence, ?e ?oul< eE)ec# #his :orm o: or-aniBa#ion #o $e more hi-hl= s)ecialiBe< in i#s a<a)#a#ion (o: course #here are #ech nical :ac#ors ?hich also con#ri$u#e #o #his as ?ell). 4ra:#-a<minis#ere< cons#ruc#ion :irms are se# u) in such a ?a= #ha# #he= can a<a)# ra)i<l= #o chan-es in <eman<, an< #he= can a<a)# #o <i: :eren# cons#ruc#ion )ro$lems $= ;ar=in- #he miE o: s+ills re)resen#e< in #heir ?or+ :orce. 8ureaucra#icall= a<minis#ere< cons#ruc#ion :irms are more s)ecialiBe< an< as a resul# #he= are e::icien# onl= ?hen <eman< is hi-h, an< ;er= ine::icien# ?hen i# is lo?. Fe also $elie;e #ha# #he= #en< #o $e more s)ecialiBe< ?i#h re-ar< #o #=)e o: cons#ruc#ion. 4ra:#-a<min is#ere< or-aniBa#ions sacri:ice e::icien# eE)loi#a#ion o: #heir niche :or :leE i$ili#=. 8ureaucra#ic or-aniBa#ions choose #he o))osi#e s#ra#e-=. This :ormu la#ion is an eE#ension o: "#inchcom$e>s an< ser;es #o sho? #ha# his ar-u men# is essen#iall= ecolo-ical. VII. DI"4%""I7C

7ur aim in #his )a)er has $een #o mo;e #o?ar< an a))lica#ion o: mo<ern )o)ula#ion ecolo-= #heor= #o #he s#u<= o: or-aniBa#ion-en;ironmen# rela #ions. For us, #he cen#ral Gues#ion is, ?h= are #here so man= +in<s o: or-aniBa#ions. 5hrasin- #he Gues#ion in #his ?a= o)ens #he )ossi$ili#=

o: a))l=in- a rich ;arie#= o: :ormal mo<els #o #he anal=sis o: #he e::ec#s o: en;ironmen#al ;aria#ions on or-aniBa#ional s#ruc#ure. Fe $e-in ?i#h Ha?le=>s classic :ormula#ion o: human ecolo-=. Ho? e;er, ?e reco-niBe #ha# ecolo-ical #heor= has )ro-resse< enormousl= since sociolo-is#s las# s=s#ema#icall= a))lie< i<eas :rom $ioecolo-= #o social or-aniBa#ion. Cone#heless, Ha?le=>s #heore#ical )ers)ec#i;e remains a ;er= use:ul )oin# o: <e)ar#ure. In )ar#icular ?e concen#ra#e on #he )rinci)le o: isomor)hism. This )rinci)le asser#s #ha# #here is a one-#o-one corre s)on<ence $e#?een s#ruc#ural elemen#s o: social or-aniBa#ion an< #hose uni#s #ha# me<ia#e :lo?s o: essen#ial resources in#o #he s=s#em. I# eE)lains #he ;aria#ions in or-aniBa#ional :orms in eGuili$rium. 8u# an= o$ser;e< isomor)hism can arise :rom )ur)ose:ul a<a)#a#ion o: or-aniBa#ions #o #he common cons#rain#s #he= :ace or $ecause nonisomor)hic or-aniBa#ions are selec#e< a-ains#. "urel= $o#h )rocesses are a# ?or+ in mos# social s=s#ems. Fe $elie;e #ha# #he or-aniBa#ions li#era#ure has em)hasiBe< #he :ormer #o #he eEclusion o: #he la##er. Fe sus)ec# #ha# care:ul em)irical reseaRch ?ill re;eal #ha# :or ?i<e classes o: or-aniBa#ions #here are ;er= s#ron- iner#ial )ressures on s#ruc #ure arisin- $o#h :rom in#ernal arran-emen#s (e.-., in#ernal )oli#ics) an< #he en;ironmen# (e.-., )u$lic le-i#ima#ion o: or-aniBa#ional ac#i;i#=). To claim o#her?ise is #o i-nore #he mos# o$;ious :ea#ure o: or-aniBa#ional li:e. Failin- churches <o no# $ecome re#ail s#oresI nor <o :irms #rans:orm #hemsel;es in#o churches. 6;en ?i#hin $roa< areas o: or-aniBa#ional ac #ion, such as hi-her e<uca#ion an< la$or union ac#i;i#=, #here a))ear #o $e su$s#an#ial o$s#acles #o :un<amen#al s#ruc#ural chan-e. &esearch is nee<e< on #his issue. 8u# un#il ?e see e;i<ence #o #he con#rar=, ?e ?ill con#inue #o <ou$# #ha# #he ma,or :ea#ures o: #he ?orl< o: or-aniBa#ions arise #hrou-h learnin- or a<a)#a#ion. Hi;en #hese <ou$#s, i# is im)or#an# #o eE)lore an e;olu#ionar= eE)lana#ion o: #he )rinci)le o: isomor)hism. Tha# is, ?e ?ish #o em$e< #he )rinci)le o: isomor)hism ?i#hin an eE)lici# selec#ion :rame?or+. In or<er #o a<< selec#ion )rocesses ?e )ro)ose a com)e#i#ion #heor= usin- 'o#+a-Vol#erra mo<els. This #heor= relies on -ro?#h mo<els #ha# a))ear sui#a$le :or re)resen#in- $o#h or-aniBa#ional <e;elo)men# an< #he -ro?#h o: )o)ula#ions o: or-aniBa#ions. &ecen# ?or+ $= $ioecolo-is#s on 'o#+a-Vol#erra s=s#ems =iel<s )ro)osi#ions #ha# ha;e imme<ia#e rele;ance :or #he s#u<= o: or-aniBa#ion-en;ironmen# rela#ions. These resul#s con cern #he e::ec#s o: chan-es in #he num$er an< miE#ure o: cons#rain#s u)on s=s#ems ?i#h re-ar< #o #he u))er $oun< o: #he <i;ersi#= o: :orms o: or-aniBa#ion. Fe )ro)ose #ha# such )ro)osi#ions can $e #es#e< $= eEam inin- #he im)ac# o: ;arie#ies o: s#a#e re-ula#ion $o#h on siBe <is#ri$u#ions an< on #he <i;ersi#= o: or-aniBa#ional :orms ?i#hin $roa<l= <e:ine< areas o: ac#i;i#= (e.-., me<ical care, hi-her e<uca#ion, an< ne?s)a)er )u$lishin-).

3 more im)or#an# eE#ension o: Ha?le=>s ?or+ in#ro<uces <=namic con si<era#ions. The :un<amen#al issue here concerns #he meanin- o: isomor )hism in si#ua#ions in ?hich #he en;ironmen# #o ?hich uni#s are a<a)#e< is chan-in- an< uncer#ain. "houl< Mra#ionalM or-aniBa#ions a##em)# #o <e ;elo) s)ecialiBe< isomor)hic s#ruc#ural rela#ions ?i#h one o: #he )ossi$le en;ironmen#al s#a#es. 7r shoul< #he= a<o)# a more )las#ic s#ra#e-= an< ins#i#u#e more -eneraliBe< s#ruc#ural :ea#ures. The isomor)hism )rinci)le <oes no# s)ea+ #o #hese issues. Fe su--es# #ha# #he concre#e im)lica#ion o: -eneralism :or or-aniBa #ions is #he accumula#ion an< re#en#ion o: ;arie#ies o: eEcess ca)aci#=. To re#ain #he :leEi$ili#= o: s#ruc#ure reGuire< :or a<a)#a#ion #o <i::eren# en;ironmen#al ou#comes reGuires #ha# some ca)aci#ies $e hel< in reser;e an< no# commi##e< #o ac#ion. Heneralis#s ?ill al?a=s $e ou#)er:orme< $= s)ecialis#s ?ho, ?i#h #he same le;els o: resources, ha))en #o ha;e hi# u)on #heir o)#imal en;ironmen#. 4onseGuen#l=, in an= cross-sec#ion #he -eneralis#s ?ill a))ear ine::icien# $ecause eEcess ca)aci#= ?ill o:#en $e ,u<-e< ?as#e. Cone#heless, or-aniBa#ional slac+ is a )er;asi;e :ea#ure o: man= #=)es o: or-aniBa#ions. The Gues#ion #hen arises( ?ha# #=)es o: en;ironmen#s :a;or -eneralis#s. 3ns?erin- #his Gues#ion com)rehensi;el= #a+es one a lon- ?a= #o?ar< un<ers#an<in- #he <=namic o: or-aniBa#ion en;ironmen# rela#ions. Fe $e-in a<<ressin- #his Gues#ion in #he su--es#i;e :rame?or+ o: 'e;ins>s (19 2, 19 8) :i#ness-se# #heor=. This is one o: a class o: recen# #heories #ha# rela#es #he na#ure o: en;ironmen#al uncer#ain#= #o o)#imal le;els o: s#ruc#ural s)ecialism. 'e;ins ar-ues #ha# alon- ?i#h uncer#ain#= one mus# consi<er #he -rain o: #he en;ironmen# or #he lum)iness o: en;ironmen#al ou#comes. The #heor= in<ica#es #ha# s)ecialism is al?a=s :a;ore< in s#a$le or cer#ain en;ironmen#s. This is no sur)rise. 8u# con #rar= #o #he ;ie? ?i<el= hel< in #he or-aniBa#ions li#era#ure, #he #heor= also in<ica#es #ha# -eneralism is no# al?a=s o)#imal in uncer#ain en;iron men#s. Fhen #he en;ironmen# shi:#s uncer#ainl= amon- s#a#es #ha# )lace ;er= <i::eren# <eman<s on #he or-aniBa#ion, an< #he <ura#ion o: en;iron men#al s#a#es is shor# rela#i;e #o #he li:e o: #he or-aniBa#ion (;aria#ion is :ine--raine<), )o)ula#ions o: or-aniBa#ions #ha# s)ecialiBe ?ill $e :a ;ore< o;er #hose #ha# -eneraliBe. This is $ecause or-aniBa#ions #ha# a# #em)# #o a<a)# #o each en;ironmen#al ou#come ?ill s)en< mos# o: #heir #ime a<,us#in- s#ruc#ure an< ;er= li##le #ime in or-aniBa#ional ac#ion <irec#e< a# o#her en<s. "#a#e< in #hese #erms, #he )ro)osi#ion a))ears o$;ious. Ho?e;er, ?hen one rea<s #he li#era#ure on or-aniBa#ion-en;ironmen# rela#ions, one :in<s #ha# i# ?as no# so o$;ious. Mos# im)or#an#, #he )ro)osi#ion :ollo?s :rom a sim)le eE)lici# mo<el #ha# has #he ca)aci#= #o uni:= a ?i<e ;arie#= o: )ro)osi#ions rela#in- en;ironmen#al ;aria#ions #o or-aniBa#ional s#ruc#ure.

Fe ha;e i<en#i:ie< some o: #he lea<in- conce)#ual an< me#ho<olo-ical o$s#acles #o a))l=in- )o)ula#ion ecolo-= mo<els #o #he s#u<= o: or-ani Ba#ion-en;ironmen# rela#ions. Fe )oin#e< #o <i::erences $e#?een human an< nonhuman social or-aniBa#ion in #erms o: mechanisms o: s#ruc#ural in;ariance an< s#ruc#ural chan-e, associa#e< )ro$lems o: <elimi#in- )o)u la#ions o: or-aniBa#ions, an< <i::icul#ies in <e:inin- :i#ness :or )o)ula#ions o: eE)an<a$le uni#s. In each case ?e ha;e merel= s+e#che< #he issues an< )ro)ose< shor#-run sim)li:ica#ions ?hich ?oul< :acili#a#e #he a))lica#ion o: eEis#in- mo<els. 4learl=, each issue <eser;es care:ul scru#in=. 3# #he momen# ?e are :rus#ra#e< a# leas# as much $= #he lac+ o: em)irical in:orma#ion on ra#es o: selec#ion in )o)ula#ions o: or-aniBa#ions as $= #he unresol;e< issues ,us# men#ione<. 4ensus <a#a are )resen#e< in a manner #ha# ren<ers #he calcula#ion o: :ailure ra#es im)ossi$leI an< li##le lon-i#u<inal research on )o)ula#ions o: or-aniBa#ions has $een re )or#e<. Fe <o, ho?e;er, ha;e some in:orma#ion on ra#es o: selec#ion. Fe +no?, :or eEam)le, #ha# :ailure ra#es :or small $usinesses are hi-h. 8= recen# es#ima#es u)?ar<s o: 45&)6 o: small $usiness :irms in #he %ni#e<

"#a#es :ail each =ear (Hollan<er 19 7I 8ol#on 1971I see also 4hurchill 1955) . In )ar# #his hi-h :ailure ra#e re:lec#s ?ha# "#inchcom$e (19 5) calle< #he lia$ili#= o: ne?ness. Man= ne? or-aniBa#ions a##em)# #o en#er niches #ha# ha;e alrea<= $een :ille< $= or-aniBa#ions #ha# ha;e amasse< social, economic, an< )oli#ical resources #ha# ma+e #hem <i::icul# #o <islo<-e. It is im)or#an# #o <e#ermine ?he#her #here is an= selec#i;e <isa<;an#a-e o: smallness no# o: ne?ness. Fe <ou$# #ha# man= rea<ers ?ill <is)u#e #he con#en#ion #ha# :ailure ra#es are hi-h :or ne? an<*or small or-aniBa#ions. Ho?e;er, much o: #he sociolo-ical li#era#ure an< ;ir#uall= all o: #he cri#ical li#era#ure on lar-e or-aniBa#ions #aci#l= acce)#s #he ;ie? #ha# such or-aniBa#ions are no# su$,ec# #o s#ron- selec#ion )ressures. Fhile ?e <o no# =e# ha;e #he em )irical <a#a #o ,u<-e #his h=)o#hesis, ?e can ma+e se;eral commen#s. Firs#, ?e <o no# <is)u#e #ha# #he lar-es# or-aniBa#ions in<i;i<uall= an< collec#i;el= eEercise s#ron- <ominance o;er mos# o: #he or-aniBa#ions #ha# cons#i#u#e #heir en;ironmen#s. 8u# i# <oes no# :ollo? :rom #he o$ser ;a#ion #ha# such or-aniBa#ions are s#ron- in an=one )erio< #ha# #he= ?ill $e s#ron- in e;er= )erio<. Thus, i# is in#eres#in- #o +no? ho? :irml= em$e<<e< are #he lar-es# an< mos# )o?er:ul or-aniBa#ions. 4onsi<er #he so-calle< For#une 500, #he lar-es# )u$licl= o?ne< in<us#rial :irms in #he %ni#e< "#a#es. Fe con#ras#e< #he lis#s :or 1955 an< 1975 (a<,us#in- :or )ure name chan-es). 7: #hose on #he lis# in 1955, onl= 2 8 (52. 1) ?ere s#ill lis#e< in 1975. 7ne hun<re< #?en#=-#?o ha< <isa))eare< #hrou-h mer-er, 109 ha< sli))e< o:: #he M500,M an< one (a :irm s)ecialiBin- in 4u$an su-arO) ha< $een liGui<a#e<. The num$er ?hose rela#i;e sales

-ro?#h cause< #hem #o $e <ro))e< :rom #he lis# is Gui#e im)ressi;e in #ha# #he lar-e num$er o: mer-ers ha< o)ene< man= slo#s on #he lis#. "o ?e see #ha#, ?hereas ac#ual liGui<a#ion ?as rare :or #he lar-es# in<us#rial :irms in #he %ni#e< "#a#es o;er a 20-=ear )erio<, #here ?as a -oo< <eal o: ;ola#ili#= ?i#h re-ar< #o )osi#ion in #his )seu<o<ominance s#ruc#ure $ecause o: $o#h mer-ers an< sli))in- sales.5 "econ<, #he choice o: #ime )ers)ec#i;e is im)or#an#. 6;en #he lar-es# an< mos# )o?er:ul or-aniBa#ions :ail #o sur;i;e o;er lon- )erio<s. For eEam)le, o: #he #housan<s o: :irms in $usiness in #he %ni#e< "#a#es <urin#he &e;olu#ion, onl= 12 sur;i;e as au#onomous :irms an< se;en as reconiBa$le <i;isions o: :irms #7ation)s Business 197 ). 5resuma$l= one

nee<s a lon-er #ime )ers)ec#i;e #o s#u<= #he )o)ula#ion ecolo-= o: #he lar-es# an< mos# <ominan# or-aniBa#ions. Thir<, s#u<=in- small or-aniBa#ions is no# such a $a< i<ea. The socio lo-ical li#era#ure has concen#ra#e< on #he lar-es# or-aniBa#ions :or o$;ious <esi-n reasons. 8u#, i: iner#ial )ressures on cer#ain as)ec#s o: s#ruc#ure are s#ron- enou-h, in#ense selec#ion amon- small or-aniBa#ions ma= -rea#l= cons#rain #he ;arie#= o$ser;a$le amon- lar-e or-aniBa#ions. 3# leas# some elemen#s o: s#ruc#ure chan-e ?i#h siBe (as ?e ar-ue< in "ec#ion III) an< #he )ressure #o?ar< iner#ia shoul< no# $e o;erem)hasiBe<. Cone#heless ?e see much ;alue in s#u<ies o: #he or-aniBa#ional li:e c=cle #ha# ?oul< in:orm us as #o ?hich as)ec#s o: s#ruc#ure -e# loc+e< in <urin- ?hich )hases o: #he c=cle. For eEam)le, ?e con,ec#ure #ha# a cri#ical )erio< is #ha# <urin- ?hich #he or-aniBa#ion -ro?s $e=on< #he con#rol o: a sin-le o?ner*mana-er. 3# #his #ime #he manner in ?hich au#hori#= is <ele-a#e<, if a# all, seems li+el= #o ha;e a las#in- im)ac# on or-aniBa#ional s#ruc#ure. This is #he )erio< <urin- ?hich an or-aniBa#ion $ecomes less an eE#ension o: one or a :e? <ominan# in<i;i<uals an< more an or-aniBa#ion )er se ?i#h a li:e o: i#s o?n. If #he selec#ion )ressures a# #his )oin# are as in#ense as anec<o#al e;i<ence su--es#s #he= are, selec#ion mo<els ?ill )ro;e ;er= use:ul in accoun#in- :or #he ;arie#ies o: :orms amon- #he ?hole ran-e o: or-aniBa#ions. The o)#imism o: #he )re;ious )ara-ra)h shoul< $e #em)ere< $= #he realiBa#ion #ha# ?hen one eEamines #he lar-es# an< mos# <ominan# or-a niBa#ions, one is usuall= consi<erin- onl= a small num$er o: or-aniBa#ions. The smaller #he num$er, #he less use:ul are mo<els #ha# <e)en< on #he #=)e o: ran<om mechanisms #ha# un<erlie )o)ula#ion ecolo-= mo<els. Four#h, ?e mus# consi<er ?ha# one anon=mous rea<er, cau-h# u) in #he s)iri# o: our )a)er, calle< #he an#i-eu-enic ac#ions o: #he s#a#e in
12 From a# leas# some )ers)ec#i;es, mer-ers can $e ;ie?e< as chan-es in :orm. This ?ill almos# cer#ainl= $e #he case ?hen #he or-aniBa#ions mer-e< ha;e ;er= <i::eren# s#ruc#ures. These <a#a also in<ica#e a s#ron- selec#i;e a<;an#a-e :or a con-lomera#e :orm o: in<us#rial or-aniBa#ion.

192

sa;in- :irms such as 'oc+hee< :rom :ailure. This is a <rama#ic ins#ance o: #he ?a= in ?hich lar-e <ominan# or-aniBa#ions can crea#e lin+a-es ?i#h o#her lar-e an< )o?er:ul ones so as #o re<uce selec#ion )ressures. If such

mo;es are e::ec#i;e, #he= al#er #he )a##ern o: selec#ion( In our ;ie? #he selec#ion )ressure is $um)e< u) #o a hi-her le;el. "o ins#ea< o: in<i;i<ual or-aniBa#ions :ailin-, en#ire ne#?or+s :ail. The -eneral conseGuence o: a lar-e num$er o: lin+a-es o: #his sor# is an increase in #he ins#a$ili#= o: #he en#ire s=s#em ("imon 19 2, 1972I Ma= 1972), an< #here:ore ?e shoul< see $oom an< $us# c=cles o: or-aniBa#ional ou#comes. "elec#ion mo<els re#ain rele;ance, #hen, e;en ?hen #he s=s#ems o: or-aniBa#ions are #i-h#l= cou)le< (see Hannan 197 ). Finall=, some rea<ers o: earlier <ra:#s ha;e (some a))ro;in-l=, some <isa))ro;in-l=) #rea#e< our ar-umen#s as me#a)horic. This is no# ?ha# ?e in#en<. In a :un<amen#al sense all #heore#ical ac#i;i#= in;ol;es me#a )horic ac#i;i#= (al#hou-h a<mi##e<l= #he #erm Manalo-ueM comes closer #han <oes Mme#a)horM). The use o: me#a)hors or analo-ues en#ers in#o #he :ormula#ion o: "if ... #henM s#a#emen#s. For eEam)le, cer#ain molec ular -ene#ic mo<els <ra? an analo-= $e#?een DC3 sur:aces an< cr=s#al s#ruc#ures. The la##er ha;e sim)le ?ell-$eha;e< -eome#ric s#ruc#ures amena$le #o s#ron- #o)olo-ical (ma#hema#ical) anal=sis. Co one ar-ues #ha# DC3 )ro#eins are cr=s#alsI $u# #o #he eE#en# #ha# #heir sur:aces ha;e cer#ain cr=s#al-li+e )ro)er#ies, #he ma#hema#ical mo<el use< #o anal=Be cr=s#als ?ill she< li-h# on #he -ene#ic s#ruc#ure. This is, as ?e un<ers#an< i#, #he -eneral s#ra#e-= o: mo<el $uil<in-. Fe ha;e, :or eEam)le, use< resul#s #ha# rel= on #he a))lica#ion o: cer#ain lo-is#ic <i::eren#ial eGua#ions, #he 'o#+a-Vol#erra eGua#ions. Co +no?n )o)ula#ion (o: animals, or o: or-aniBa#ions) -ro?s in eEac#l= #he manner s)eci:ie< $= #his ma#hema#ic mo<el (an< #his :ac# has cause< numerous na#uralis#s #o ar-ue #ha# #he mo<el is $iolo-icall= meanin-less). Fha# #he eGua#ions <o is #o mo<el #he -ro?#h )a#h o: )o)ula#ions #ha# eEis# on :ini#e resources in a close< s=s#em (?here )o)ula#ion -ro?#h in #he a$sence o: com)e#i#ion is lo-is#ic an< #he )resence o: com)e#in- )o)u la#ions lo?ers carr=in- ca)aci#ies in #ha# s=s#em). To #he eE#en# #ha# #he in#erac#ions o: )o)ula#ions o: 8aramecium aureilia an< 8. caudatum (Hause>s eE)erimen#) mee# #he con<i#ions o: #he mo<el, #he mo<el eE )lains cer#ain +e= :ea#ures o: )o)ula#ion <=namics an< #he rela#ionshi) o: en;ironmen#al ;aria#ions #o s#ruc#ure. To #he eE#en# #ha# #he in#erac#ions o: )o)ula#ions o: ra#ional-le-al $ureaucracies an< )o)ula#ions o: )a#ri monial $ureaucracies also mee# #he con<i#ions o: #he mo<el, #he mo<el eE)lains #he same im)or#an# )henomena. Cei#her #he )ro#oBoa nor #he $ureaucracies $eha;e eEac#l= as #he mo<el s#i)ula#es. The mo<el is an a$s#rac#ion #ha# ?ill lea< #o insi-h# ?hene;er #he s#a#e< con<i#ions are a))roEima#e<. 192

Throu-hou# ?e ma+e a s#ron- con#inui#=-o:-na#ure h=)o#hesis. Fe )ro)ose #ha#, ?hene;er #he s#a#e< con<i#ions hol<, #he mo<els lea< #o ;alu a$le insi-h#s re-ar<less o: ?he#her #he )o)ula#ions un<er s#u<= are com )ose< o: )ro#oBoans or or-aniBa#ions. Fe <o no# ar-ue Mme#a)horicall=.M Tha# is, ?e <o not ar-ue as :ollo?s( an em)irical re-ulari#= is :oun< #o hol<

:or cer#ain )ro#oBoansI $ecause ?e h=)o#hesiBe #ha# )o)ula#ions o: or-a niBa#ions are li+e )o)ula#ions o: )ro#oBoans in essen#ial ?a=s, ?e )ro)ose #ha# #he -eneraliBa#ions <eri;e< :rom #he la##er ?ill hol< :or or-aniBa #ions as ?ell. This is #he +in< o: reasonin- $= ?hich $iolo-ical )ro)osi #ions ha;e mos# o:#en en#ere< sociolo-ical ar-umen#s (e.-., #he :amous or in:amous-or-anismic analo-= a<;ance< $= ")encer). Ins#ea< o: a))l=in- $iolo-ical la?s #o human social or-aniBa#ion, ?e a<;oca#e #he a))lica#ion o: )o)ula#ion ecolo-= #heories. 3s ?e ha;e in<i ca#e< a# a num$er o: )oin#s, #hese #heories are Gui#e -eneral an< mus# $e mo<i:ie< :or an= concre#e a))lica#ion (sociolo-ical or $iolo-ical). 7ur )ur)ose has $een #?o:ol<. Firs#, ?e s+e#che< some o: #he al#era#ions in )ers)ec#i;e reGuire< if )o)ula#ion ecolo-= #heories are #o $e a))lie< #o #he s#u<= o: or-aniBa#ions. "econ<, ?e ?ishe< #o s#imula#e a reo)enino: #he lines o: communica#ion $e#?een sociolo-= an< ecolo-=. I# is ironic #ha# Ha?le=>s (19!!, ). 299) <ia-nosis o: some 20 =ears a-o remains a)# #o<a=( M5ro$a$l= mos# o: #he <i::icul#ies ?hich $ese# human ecolo-= ma= $e #race< #o #he isola#ion o: #he su$,ec# :rom #he mains#ream o: ecolo-ical #hou-h#.M
&6F6&6C46" 3l<rich, Ho?ar< 6., an< Je::re= 5:e::er. 197 . M6n;ironmen#s o: 7r-aniBa#ions.M Annual 9eview of Sociology 2( 79-105. 3l<rich, Ho?ar< 6., an< 3l$er# J. &eiss. 197 . M4on#inui#ies in #he "#u<= o: 6co lo-ical "uccession( 4han-es in #he &ace 4om)osi#ion o: Cei-h$orhoo<s an< Their 8usinesses.M American Journal of Sociology 81 (Januar=)( 8! - . 8lau, 5e#er M. 1972. MIn#er<e)en<ence an< Hierarch= in 7r-aniBa#ions.M Social Science 9esearch 1 (3)ril)( 1-2!. 8lau, 5e#er M., an< &ichar< 3. "choenherr. 1971. :he Structure of ;rgani<ations. Ce? 9or+( 8asic. 8lau, 5e#er M., an< F. &ichar< "co##. 19 2. =ormal ;rgani<ations. "an Francisco( 4han<ler. 8ol#on, J. 6. 1971. Small =irms. &e)or# o: #he 4ommi##ee o: InGuir= on "mall Firms. 'on<on( Her Ma,es#=>s "#a#ioner= 7::ice. 8oul<in-, Nenne#h. 1952. MTo?ar< a Heneral Theor= o: Hro?#h.M Canadian Journal of >conomics and 8olitical Science &5?2'@$A6. 8urns, Tom, an< H. M. "#al+er. 19 1. :he ,anagement of *nnovation. 'on<on( Ta;is#oc+. 4a)lo?, Theo<ore. 1957. M7r-aniBa#ional "iBe.M Administrative Science Buarterly 1 (March)( !8!-505. 4hurchill, 8e##= 4. 1955. M3-e an< 'i:e 6E)ec#anc= o: 8usiness Firms.M Survey of Current Business 25 (Decem$er)( 15-19. 4roBier, Michel. 19 !. :he Bureaucratic 8henomenon. 4hica-o( %ni;ersi#= o: 4hica-o 5ress.

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