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Quantitative Genetics

Quantitative Genetics

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Published by: holacanthus22012 on Aug 21, 2009
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Anyone who has sampled populations of fishes will agree that tliere is great phenotypicvariation in fishes. Conspicuous and economically important differences in growth rate,bodysize at maturity, and other traits frequently occur within and between populationsof many fish species (Allendorf et al. 1987;Table 12.1).Although morphological charac-ters cannot be compared directly among taxa, fish are phenotypically more variable thanare other vertebrates (Mayr 1969:170). Coefficients of variation among individuals
populations (Table 12.1, upper panel) are typically less than 10% in other verte-brates but exceed that leve1 for most morphological characters in fishes.Variation
conspecific populations of fishes (Table 12.1, lower panel) is generally larger than that inother vertebrates, often severa1 times that among species in other vertebrate taxa. Suchwide variation in quantitative characters raises a number of questions that fisheries sci-ence must strive to answer. How much of the observed variation is due to genetic fac-tors? What is the adaptive and evolutionary significance of this variation? How shouldwe consider quantitative genetic variation i~iisheries management and cotiservationactions?This chapter on quantitative variation might have been presented in the first sectionof this text along with those on molecular and chro~iioso~nalariation. However,
elected to defer discussion of quantitative genetics until tlie reader became familiar withpopulation genetic processes. The intent of this cliapter is to provide a foundation forunderstanding how quantitative variation is nieasured and its genetic basis inferred andthen to discuss the implications of quantitative genetic variation for fisheries manage-ment and conservation.
Recognizing the Nature of Quantitative Traits
Quantitative traits
exhibit a range of phenotypes that are measured as opposed toscored into a few discrete classes. Quantitative traits iticlude such adaptively importanttraits as morphological, physiological, and behavioral characters.There are three generalclasses ofquantitative traits, distinguished on the basis of the distribution of phenotypes.
Phenotypic variation within and between populations for chosen characters in fishes.Adapted from Allendorf et al.
Variation within populationsCoefficient ofSpecies Character variation (CV)" SoiirceRainbow trout Body length (140 d-2.5 years)10-19 Gjedrem (1983)
Onrorhynchus mykiss
Body weight (150 d-2.5 years) 17-56Atlantic salmon Uody lengtli (190 d-3.5 years) 7-23 Gjedrem (1983)
Salmo salar
Body weight (190 d-3.5 years) 25-76Common carp Body weight (adults) 22 Gjedrem (1983)
Cyprinus carpio
Channel catfish Rody weight (juveniles) 46 Gjedrem
Irtalurus punctatus
Body weight (adults) 27GLIPPY Body length (28-63 d) 10-1 2 RymC1ri
Porcilia rrticulata
Body weight (42-63 d) 35-36Variation between populationsLargestSmallest value as
Species group Character measurement of smallest SourceArctic char
Salvrlinirs alpinus
5 Palearctic stocks) Body length at 2 years 25 mm 800 Johnson (1980)Body length at 4 years 80 nlm 398Body length at 10 years 145 mm 417(10 nonmigratory Weight at maturity 23 g 4,213 Johnson (1980)Palearctic stocks) Number of eggs 48 6,465Urown trout Body lerigth at 3 years 11.1 cm 270 Alm (1939)
Salmo trutta
Body lerigth at 6 years 19.3 cm 337(1 1 European populations)European perch Body length 9.8 cm 315 Alm (1946)
Perca Juviatilis
(23 Swedish populatioris)Lake whitefish Number of gil1 rakers 23 150 Lindsey
Corrgonus clupc~~fi~rmis
(2 synipatric forms)
(staridard deviation/rneari)
Continuous traits exhibit a continuum of phenotypes that are measured. There areinfinitely many posible phenotypcs, among which discrimination is li~nited nly by ourability to nleasure them. Examples of continuous traits include growth rate, yield, andmorphonietric traits (e.g., head 1ength:standard length, eye diameter:head leiigth, andbody depthxtandard length).Meristic traits exhibit phenotypes expresied in discrete elements that are counted toform integral classes. Ari individual's phenotype is characterized as the number of ele-inents of the trait it exhibits. Examples of meristic traits in fishes are number of scales
Quantitative Genetics
along the lateral line, number of fin rays, and number of pyloric caecae. When the num-ber of elements exhibited becomes very large (e.g., fecundity), the distinction betweencontinuous and meristic traits vanishes.Threshold traits are discrete traits that are either present or absent in an individual.Generally, both genetic and environmental factors affect expression of the trait. Observa-tion of the trait in a particular individual implies that the individual has a liability overthe threshold for the trait's expression. This liability is not always directly observable; ageneticist would need to study an individual's pedigree to infer the likelihood that thisindividual may express tlze trait.The best-known and studied threshold traits are thehuman diseases diabetes, sclzizophrenia, and certain cancers. Exemplifying the geneticand nongenetic factors affecting expression of a threshold trait, the likelihood that a per-son with a genetic predisposition for diabetes will actually suffer the disease will alsodepend upon his or her diet, weiglzt, and age.Threshold traits also are observed in fishes.Geneticists hypothesize that the tendencia of certain salmonids to undergo precociousmaturation and to pursue anadromy are threshold traits.
Recognizing Qualities Common to Al1 Classes of Quantitative Traits
It is not possible to infer from phenotype the action of segregation and single-locus-mediated gene expression because expression of quantitative traits can be controlled bymany genes, a property termed polygenic trait deternzination.Tlze infinitesimal modelfor expression of quantitative traits assumes a large nunzber of loci eaclz contributing asmall and equal amount to expression of the trait. In reality, however, the contribution ofindividual loci to expression of the trait rangesfronz snzall to large. Should one gene playa large role in determining expression of a quantitative trait, it is termed a major gene.For example, rnajor genes are known to affect expression of double muscling in BelgianBlue cattle and fertility in Beroola sheep and Meishan pigs.The nuniber of genes affect-ing expression of certain quantitative traits has been estimated (Box 12.1).A particulargene may affect multiple traits, a condition referred to as pleiotropy. Because many locigenerally are involved in the expression of a quantitative trait, al1 of which segregateindependently (excepting linked loci), a large range of genetic variation is common.Quantitative loci exhibit the same niodes of inheritance as Mendelian, or qualitative,traits (Chapter 1) except that they are influenced by alleles at many loci. Each locus seg-regates in accordance to Mendelian expectation. Following meiosis (in a diploid organ-ism), each gamete contains one of tlze two alleles carried by the parent. Alleles at a locusinteract with each other according to Mendelian modes of phenotypic expressiondescribed in Chapter
Expression of quantitative traits is affected by environmental (i.e., nongenetic) fac-tors. Sensitivity to environnzental effects will differ among loci. Polygenic trait determi-nation and environmental influence give rise to continuous distributions for quantita-tive traits.Because phenotypes are the consequence of multiple, interacting genetic and envi-ronmental effects, it is not possible to determine an individual's genotype from its exter-nal appearance. However, quantitative geneticists have developed a rich body of theoryon how to apply statistical inference to quantify genetic and nongenetic factors underly-ing expression of quantitative traits.This body of theory is surveyed in the next section,preparing us for discussion of the implications of quantitative variation for adaptationand evolution.

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