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Con~ewationis charged with thepresetyation of a dynamic, ever.

changfngsystem of vanafroncontained in each species.
Hampton L. Carson
Modern eenetics (the stu~ d vof hereditv and variation) has reached into
old-fa\l:ioned concepts. One set
every corner oi biology, d r a s ~ i & l lchanging
of new discoveries has to do with the existence of generic variation from
The natural pwu!ation of the
individual to individual in natural ~~lpulationr.
human species displays suchvariabilliy, and it is not hard foithe casual
observer to recognize that both genes and environment are involved. Some
domestic animals (for example, dogs and cattle, Canisfamiliari.7, Bas taurus)
also show easilv observed individual neneticvariation. This is an old storv:
inf&mation frjm the n& &eneri&eClend.:bur perception and indicates (hat
nearly every normal, sexually reproducin~species is re lete with genetic
vartntion. much of it st the molecular levcl. Alrhourli [idden from easv ohbervntion. permanetit molecular variation from individh to individual is i o allpewasive throughout nature as to constit~tean nlmoht un:\'ersal law of life.
In the last Century, wild animals and plants were frequent2 viewe? as if
the individual members of a species were basically identtcal. ach mdividual
was expected to conform to some kind of type; any deviation from the type
was considered somehow unusual or accidental. The idea of twes is reallv a
human concoction used to simpl~fythe way we deal with natuii. Fxcept pkrhaps as a device for cataloging material in museums, we mutt ab:indon the
conccpt of the type, since gcneticq has shown it to he bad biology. We now
know that not only is each human individual unique both in genetics and
environment, but so is each dog or elephant or oak tree.
It is imoortant to recoeniz; that the conservationist is not chareed with
the prese&ation of types. Indeed, what is out there in nature is a dynamic,
ever-cliangin~sptem of variation contained in each species. It is this
svstem that lies at the heart of each of the oreanisms that
flexible vGiaiiin -wc study or va!uc ss something to be preserved. As contempo;ary biologist\,
u e arc view~ngthe systcm at only one moment in the progre.;? of geolopcal
time. The co6servaiionist must iraoole with the difficultidea of ihe nonpermanent n m r e of horh the sp'kciei and the ecosystem in which it exists.
Although some species and ecosystems are more stable than others, ewlut i o n a ~ ~ c h a wlth
n ~ e time is alaw of life.
Gkneticiariation arises through continual, irreparable, mutational
changes in the D ? A carried in the cells. Most of these mutations have on1 a
smal effect indtv~duallv:the eenetic uniaueness of individual.; is eenerarc (I"
genetic recombination: i h i s 7s the naturally occurring ~crambling~rocess

tI1;tt the genes undergo each time the sex cells, egg and sperm, arc produced
and then combined to form a new individual. Genetic recomhinat~onis a
nearly universal accompaniment of sexual reproduction. which -is the renro~
ductive mode of almosi all of the forms of lifk that we- ----,
mav un.ch
to m .n&
-..G .
Some species, particularly among plants, have developed a capacity for clonal
reproduction, in which a little piece of the mother plant becomes seoarated.
as, for exam le, in a propagated cutting., The pie& can be induced i o grow'
into a new p ant that is truly genetically den tical (except for new mutations)
to the plant from which it was taken. A single mother plant can be used to
make thousands of such cuttings. When planted out in afield.
the r ~ w l t.i.n o
- - - .--- .plants may ]oak like a population, hut t h i colleclion does not have the varr
ab~lityof a population; ~tis re:dlv only one 1n(l1\,1dua1.
Cloning. procedures
are wholly impractical as a means for conserving gene nook: th& can
merely perpeiuate one or a few combinations of genes; mo&fiedon& slightly
by the accumulation of non-recombining new mutations.

-- --


The discoven, of extensive genetic variation in natural ~ooulations
, ,
~ led
early pop~~lation
genelici.;tr to &fer to the gcrictic m:lterial carrled by the
individuals of the population a< a 'pool i r f genes. Ihic term is used to suge
a so? of collective hereditam,endcw-zest the fact that a t ~ uo~ulation-has
men1 that hclongs to thi bieeding group. In a sense, each time reproduction
occurs, the individuals pool their genes i n the mathematical sense. It is
derives its genetic
from this pool that the next gcncrarim I I intli\.iduals
endowment. Ai long as the population of a zpecies remains large, natural
selection will tend to preserve gcnctic variahilit) :n the gene pool.



Contram to earlier views. variabilitv in the eene 0001 does not consist of
just :I small ;lumber of variabie genes of major gffect'scattered within a mostly
or homozygous, hackground. In contrast, the heterozygous state
is the rule almost throughout the DNA. Most of the effects of these genes in
hnth natural and artificial populations, furthermore, are individually small.
Thic does not mean that they lack importance, however. Early in this century
i t was ertahlishcd that most of the adantivelv imnortant characters of hoth
plant, and animals (called the quantitbtivefharierers) are polygenically
mean~ngthat many aenes of individually small effect contr~bute
to each im~nrtantAaracter. 'Thus. there is an additive as well as an interactive som'poncr~t,and each character is affected bv many genes. This important f:wt ha\ heen somewhat concealed by the preoccupation of the human
mcdlc;tl genetic~stwith genes of large effect. These,are mostly serious
acciJcnr;d mutations, amountin to defects in funct~onor structure; almost
ey are not the raw matcnal that natural
a11 are pathological in nature.
selection used to huild the human genotype. 'lhc importance ofthe small
mutation was the original discoverv of the experimentd animal and plant
breeders, who demonstrated the effects of artificial (man-induced) selection
through their studies of the inheritance of quantitative characters. Natural
selection is not different from artif~cialselection in itc basic attrihutes.


Gene Pool Conservnliott \Carson


Accordingly, the hreeding process in both natural and art~iicialpopulartons

tends to Dreserve complexgcne comhinations that serve the needs of the
nr~anism(orthe ~raciicalbreeder) best in the heterozveous state.
At r e p m d u c t h , the exist in^ gene comhinations aiereshuffled. and novel
combinanons. perhaps only slightly difierent from thcir progenitors, are
eenerated. These genetic refinements are continuallv being tested in natural
populat:ons. Con&rsely, recombinations prod~.cingblocks-oi gene. in the
hnmozvgous state tend to have a lower fitness. so that population s u ~ v a l
comes to depend more and more on individuals that are comolex multieenic
heterozygotks. The result is what is called balanced genetic ~ l y m o r p h h n ,
a cond;tion [hat ?,sentiall) assures the mainten:lnce of eutcnsiw genetic
variabilitv. At the same time. however. the svstem exacts a ~rice.sincethe
individuds in the
arc quite uneq;al in their cap~citvio produce
progeny. The price consists of the necessary production oi a number of gene
combinations that are relativeh inferior rer~roducer\.Scw laree or conwicuous mutational changcs are in most case\'deleterious and are>limmate* by
selection; or (as i n the case of industrial melanism in moths, for emmple)
they are retained in the presence of many other gencs ofsmall effedt that
have hcen selected, so that they modifv and refine the biological properties
of the main biological charaster (in this c u e , protective n~imicrv).There is
no such thing as "5ne gene, one adaptation."
The non-geneticist tends to interpret the above emphasis on heterozygosity as meaning that ,qn~and heterozygosi must be preserved at any
cost in order to kee apopulation genetically h e z h y . However, the maintenance of specific " alanced heterozygosity' which serves the needs of the
individuals in the populations is the crltical point.



When a natural population is large, balanced polymorphism is maintained by natural selection, and therc is usually m p l e opportunity for the
fine-tunine nf the eenetic bxis of both adaotation to the environment and
rcProductLe effic'kq. A, was ment,oned above, the raw marcrial. that are
incorporated are mostly genes of verv small individual effect. When the size
of a natural ~ooulationb~comes
reduced. however. the eenetic variabilitv on
which the fldxibi~it~
of adaptive properti& depends mayYhcpartly loht, o i the
organization of i t mav he dkturhcd due to chance loses.
- Reduction in ~ouulationsize constitutes a threat to the ~ o o u l a t i o nand
this is manifested in ;ever31 wap. In clas3ical population &neiics, emphasis
was placed on loss of geneticvariability due to the fact that breeding from
only a m a l l nurnher of individuals introduces a sampling error of reproduction, such that some cne5 in low frequency in the parental population may hc
h b.v chance from t i e descendant population. Such a loss is termed random
drift and ha.. often been referred to as the Sewall Wrieht
" effect after its
~. ~~.
Drift is surely an important source of variability loss, hut ,uch hottleneeking, or unrepresentative constriction, of a population also has a disturbing effect on the genic equilibrium that underlies adaptive characters, cdusing genetic balance to undergo an equilibrium shift. The smallness of the,
population provides less room for thc acc~immodationof the proce\s of trial

Gene Pool Coflservation\Camon


recombination. The system of adaptation is blind to what may be needed in

the environment, so that when the population's ability to generate lots of
variants is impaired, this also adversely affects adaptive response. In brief,
a good adaptwe response and efficient reproduction of a population depend
on the maintenance of balanced genetic polymorphisms. This balance 1s upset in populations that have been reduced to a small size.
Following a population bottleneck, several outcomes are possible. The
first, and the one that concerns the conservationist the most, is that the population may be threatened with extinction. Some of the genetic details that
underlie the death of a population or species are well modeled in what happens in experimental populations and will be considered below. Basically,
there is a loss of both genetic variability and its organization. If the reduction in p<~pulatiun
si&is nioderm, th& m:lv be &rely a shift from 3n older
halanced condit:on 111 an dtered one. The nesr hdlance m3y serve the popul;~.
tion equally uell. or i t inay pcrmit the urganim to better meet a ncw ecologic;tl challenge that might
been instrumer~talin the population decline. In
such a c;ise. the genetics o i t l ~ cpopulation c h a n ~ e sto rncr.1 the changed cun~lit i a m fulnu,~ng
the hottleneck. Such genetic shifts mov ucc.lr rclativcly
quicklv hut are not very diifcrent from .imil:~r b;~lt,ilceshift>that occur i n
larger pop~.liltions. I n sumc csset, howe\,er, the altered population mnv shou
new conspicuou\ charxters (ior extmple, size, color of hmr) that reflect the
underlying genetic shift. Indeed, s thcury of species forniat~tmhas heen dewluped around this ~~hcnomcnl~n
of genetic reorg3nizatinn b recomhins~i~m.
In mnmaty, drastic red~~ctic~n
i n populxiun sile nit,) be olloncd by:
I ) an apparent nm;~inten:~rii~~
uf the .srurrr.\qiro; 2) 4 wdrlcn ~ h i i in
t impurtant chttracrer,; or 31 cwtincti~m.Extinction is ccrt:~inlvh\ idr the must
probable outcome of severe population bottlenecks.


The r.onsc.nutioni>tis pr~niarilvconcerned with popu1;ttions cxist~ndin a
.itsre of n:tturc, and t l i i b is the rearon for the cmpha~t\placed a h w e nn
natural populations. On the other hand, there a?e noGan increasing number
of instances where the sizes of the natural nooulations
of valued suecies have
, ,
hecome so low that speci:,l or e\trhurdin:lry measures ;,re ireq~en'tlypropth.4
for saving them. I'rumincnt : m m g these is thr. remwal of all specimens from
nature and re.wrting. at 1cad teml1~3rily.to p r o p ~ g ~ t i oinn captivitv. Examples arc the ('3lifornia condor and the 1lhnaii;tn crow ( G w ~ r n o ~ p \
T h e cxpcctcd genr.tic reil.lr< of cnptiw breeding or [ ~ r o p a g : h nare
hrietlv outlined IieIu~..The geneticist is in a reasonahl!. good pobitiun to
make judgments on such procedures, since there is an exiensive literature on
the eenetlcs of artificial oooulations of domestic animals and olants. includn~
ing &ensive experimcnial'\rork on lrlhnratnq p n ~ ~ u l a t i uu<~n,u,l)liilo
(fruit or p u m x e f l ~ e t )mice.
:mi other species. Cunsider;~hlcevidence exist>
Illat one uf the in;tjor pruhlcins ariring in m:Jl c:tpti\e populations is hreakduun in the mating s!steni,; this >criously interferes nit11 the reproductive
process. I n some birds :dinsects. mide choice (either fem;~lechoice or 1 1 1 . k
choice) depends on the e\i>tence o i a genetic;~ll! v;tri:~hlefield ui indi\.idu;lls
that 3re suhject to the t)stein oiclioice. Such iid& of mating clmice

Gene Pod CONm6on IGuson


naturallv tend to be large (for examole, mating swarms of male flies such as
midge,): if reduced in &e, repruduitive etiici:ncy appears to iall off.
When captive populations are very small, the tendency exists for the population manager to bubsrit~tehislher oerson:~lchoice oimates for a naturd one.
Sexnai selection. ooeratide thrbueh mate choice. is a oowerful svstem
for efficient reproducti'on t h a t y e ~ i yo
s maintain genetic halance, hiternzygositl, and wgor in populations. In some expcr~mentalpopulations of
inxects opert~tingunder sexual vAection, one-third of the males arc far more
suecedul in reproducing than the other two-thirds. Were the opulation to
he reduced by chmce to just a few males from the less succes.;t!l
cnte ow a
subsequent decline in vigor of the populathn wnuld iollow. Choicc oka il\gllfitnesr male from a field ofgenetically vari3blc males is strongly indicated
in i h e x caws. l i t h e iield nimdes from uhich to choosc is spsrse, the
female mav eo unmated even though what amears to the obselver as "eood"
matlngs in such ca,es also
males ;ire br&ent. l'hc irrqul:lr siccess of
related to the phenomenon of female choice. Success of captive breeding
programs in hirih, for example the condor and the 1 hw:,iian crow, :,re likely
to run into difficulties of mate choice.
The above ohservatiorls suggest that, in breeding program>. every effort
should he made by the managers of the populations to allow the natural cholce
sptem that has e\olved within the specie. to opcratc in the ex lerimenul
program. 'I'he females of many specles are far hettcr judges 0 the reproductive value of the different males that mav he availa5le than is the human
6 e mdte choice principle, although probdhly very w~despreadin animal.
and to some degree in ulants. shnuld not he a w m e d to lx an absolute rule
for the matine 6f all cabtive soecies. In the case of some rare unmlates in
zms, tor exmple, a crafted progani of ni:de select~mb;,;ed on known xncc,.
try mav be a necessav suhstitution for free mating choice. Such a rogram
uas ormoscd h\ Temdettm (1950) and r~~ccei~fullv
imolemented or thc m:,li.
ngemeni of the h r e e t h g rem;lant bf speke's galelk (doze//ospekei) in zoos,
where Lery ieu male. were avail:lble. Eswntisllv, the scliemr call, for rxtsitive artificial selection favorine a novel genetic &em in soecies that can
uithstmd close mating of relaikes (inb;&ling).' Such s;hkmcs can be
instituted, however, only in certain sprci't I casts.
At this noint. it i, rele\.ant 111 recmoh:i\ize that the nrincinal chanec that
is induced by a hottleneck in populatinn size i:. in
or dnizatioc
Disorpnization ir not necersarilv lethal to the population. hus, .I shlft in
genetic oreanization mav result once a oooulathi has successfuliv survived a
critical lo;; of its origind genetic orr:in;z:;tion. A rrorg:mizetl chbracter
may emerge in a few eener;~tio:is. l'l~isnl~\.elcliaracter m w thus be imlmec!
on The o o d a t i o n bv an altered orocess of selection that oderates on thk
disorg;;ni;cd genttit rcmn:lnts t;i the former orqani/;ltlon.'
A st~br~opulation
derived in this manner is frequently much less vigorous
than the okiehal one. because of the nhase of imbalance through whi& the
population must go Gelore natural seiection produces a new hdance. in very
extreme ca,cs o i popolatinn SIX red.lction, where the new popdatmn ctems
from one or vervfew foundinz individuals. it has been o r o ~ o i e dthat an active
period of d i r e c t h a l ic.lectio<may temporaril~replxe'the'former Ihalancing
selection. If the population can survive this, the result, aiter a series of gcnerations th;it a l l w thc ooriulari~~n
size to huild uo :wain. mav he mmifcsted
in a population with a
altered gene pbolr ~ n s h o i tsuch
a series




f . :

Gene Pool Conservation \ Carson


of events has been pro~osedas a mode wherebv new s~eciesevolve: however.

shifts induced in 'x~)criment;tl p ~ ~ ~ ~ u l a tare
i o nhot
s thi<ir,~found.
Artificial breeding hyslcms are attempts to suhstit~tea new gcnelic \ywtn
for the one that exided in thc nxtural ~onulation.7'hi. is a tall order. and
success in such a venture also has a
There can be a substantiailoss of
certain very desirable characters. For example, female choice among bower
birds (Ptilonorhvnchidae) or birds of ~ a r a d i s e(Paradiseidae) d e ~ e n d on
s vew
o i male;. In nature, (he forest has
elah~ratese~ondilryS P X U ~ characte&
many wni,peting malt$, caih d i q ~ l q i n gch3r;lctr.r that appeal not only to r h ~
reproduct~\et ~ e ofs the ierncile>hut s l v ~to the human observer's aethctic
sense. lJnder engineered programs of inhrrcdin~,:~ssumingthat they would
work, such I;~;c~~iating
plumage and hehs\,ioral character, ~ o u l dhe expected
to erode under artificial \ ~ h ~ i to
o na n t ~ ~ munhne
set of charmers. \Vh:,t
i\ happening eenelic:tllv i.; t l i ; ? ~lhe m;ln;tger or hrceder i h ewlving a genetic
+stem u hich he finds 'n~:~n:~ge;~hle,'
l'lic pop~~I:~tion
that ix ultini:ttely
t~re\en.cdm:n he hii~lu~ic;~llv
of the
- . far di>t;~ntirom the orizin:J
- -a~;il.i
preservation program.


some other ipe&, not strongly developed. h he mating of close relatives
seems not to be deleterious, nor has any clear loss of vigor followed inbreeding. Indeed, the breeding of close relatives may be natural to this
species, making genetic management of small populations a relatively easy
matter. Possibly, because of the history of repeated founder effects in the
evolution of Hawaiian suecies. the conservationist mav be dealing with
genetis h!htenir that h:~\hh~~ilt-in
mech;~ni.;mito p r e s k e genetic\wiahilit!,
t ~ tarc
~ > ireq~tentlywhjested to $e\ere reductionr or
evcn i n p ~ p u I : ~ t ~that
bottlenecks through a few individuals.
This t w e of oreadaotation exists in some terrestrial s~eciesnaturallv
\,;;>I arc:,; of open ocean. Some of the n a t h hcach plank
of lldndi'i, tor eumplc. :Ire adaptcd to long-disttrnce di5persal and show littlc
tendcncv 10 v a y ctrsst~c;~ll!iruni island to i\land. I-ittlc 1s known of properties of genetic h;,l:~nx,hut amne scem likelv to he adapted for the capaciry
eifectively h\pdhsiny
to reprod~ceafter arrival hv onli ;1single pr~ll~ngule,
normal outcrossed sexual reuroduction.

exce6tion rather than the rule.

It is possible to end this brief account on the following positive note.
Since most Hawaiian organisms have a long history of going through bottlenecks, there may be some degree of built-in ability to withstand the

Gene Pool Consen~a~iort\Camon


increasingly severe population bottlenecks now being imposed by widespread

environmental destruction. Verv few soecies. however. can be successfullv
pro agared in the lahoratory, f'eid
c x p e h e n t a l farm, or mo. l'hei;
ecokgical requirements are often too spectal. Only a minority of Hawaiian
Droso>hila soecies. for examole. can b e b r e d in thelaborator? even for
those'that can he so hred, iniportant natural characteristics a i e evidently
10% Shture is ;I better stockkee er than we are, and this principle would
appear to apply to many kinds oforganisrns. From the g
m& point of view,
the estahlisllment of manv rehtively small reserves ma!, serve ar viable
reiugia ior I lawaiian plants and an~mals.&e size of a successful rehgiurn
in Hawhi'i should be determined by what it takes to provide maximal insulatiun of [he ecosystem from encroachment b aliens and t o rotect it from
other adjacent degrading forces (see also d l t , this volume!.
A large number
o i refuria of ec~~loeicnllv
sound size will ~ r o b a b l vserve Hawaiian consewation needs better than <fewlarge areas.'

Important References
Carson. H.L. 1986. Patterns of inheritance. American Zoolopisl 6797-8(n.
Canon. ILL. 1987. The process wbcrcby species originate. B;nrctenre 37(10):715-720.
Cmant, 5. 198.8. Sdving cndangcrcd rpecie, hy trmslou1i.m Biorrmcc 38(4):254-257.
Frankel. O.H..
aod M.E. SoulC. 1981. Conremation and Evoi:dion. Cambridec
- Uniwrritv
press, &bridge, England.
He- J.P., and J.K Hodges (eds.). 1985. Advonces in Animal C o n s e ~ t i o n Clarendon
Press, Oxford, United Kingdom.
Holl, A. [Thisvolume] Protection of nahlral habitats.
Schanewald-Cox,C. 1985. Genelieg minimum population size, and the island prexrvc.
Pp. 432458 IN C.P. Stone and J.M. Scott (eds.),Hawoi'i's TerresrriolEeosysrems:
Premvarion andManogenmrr. Univ. Hawaii Press Tor Univ. Hawaii Cooperative National
Park Resources Studies Unit, Honolulu.
Scbonewald-Cor, C.M., S.M. Chambers B. MacBryde, and W.L. Thomas. 1983, Generics and
CmeIyalion: A Reference for Manoninn
" W l d Aninlo1 and Pbnt Pomloriom.
Benjamin/Cummings Publ. Co., Menlo Park, California.
Riolw: The Science ojScarciry andDiwmiry.
Soul4 M.E. (ed). 1986. Cow-ti01t
Sinauer Assc&tes. .Inr..Sunderland. h & a c h u ~ t t r
Templeton, A.R. 1980. A theory of speciation via the founder principle. Genetics