9
Heritability and Genetic
Advance
INTRODUCTION
deitability and geneuc advance are
important selection parameters. Heritability estimates
advance
genetic are more helptul in
predicting the
along
mates alone. However, 1t s not
gain under
selection than hentabiny
necessary that a character showing high hentability
exhibit high genetic advance (Johnson, et al.
1955). A brief wil
presented as follows. description of these estimates is
HERITABILITYY
The ratio of
gepotypic variance to total or phenotypic variance is known as heritability. It is
generally expressed in percent. Thus heritability 1s the
sa good index of the transmission of
heritahlc portian ol phenotypuc_Yariance.
Estimates of
characters from parents to.their offspring (Falconer, 1981).
heritability are helpful to the plant breeder in the selection.ot.clite.genotypes from
verse genetic populations.
TYPES OF HERITABILITY
inthe calculation of heritability cither genotypic or additive genetic variance
vanance is used as
numerator. Based on calculation, herntability 1s ot two ypes, VIZ, broad sense
rOW sense heritability.
heritability and
. Broad Sense: Heritability w
LakM
7Maa ud A
The ratio of genotypic variance to oa 0r pncnopie vaniance) 1s Known as broad sense
eritlability, It is calculated from total geneuc vanance which consists oft additive, dominance and
static variance. The main features ot broad sense heritability are presented below
Kxenilas of Plant
Brvedy
useful in animal breeding than plant breeding It i
I. Application : It is more
material,
selection of elilte genotypes
from homozygous
well as
calculated from both parental
as
segregating
Total
2. Calculation
genetic
: It can
variance is used as
be
numerator in the calculation population
po
3. Formula Used: It is calculated with the help of Tolowing Tormula (Manson et al, i
1996
Heritability (h8) Vg/Vp Vg/(Vg+ Ve), where Vg, Vp and ve are genotypie, phenon
and environmental variances, respectively, The estimates or heritability are rated as low (le
(len thas
S0%) and very high (above 806),
406), medium
(40 to 600), high (61 to
B. Narrow Sense Heritability
The ratio of additive genetic variance to total or phenotypic varance 1s referred to as namoy
sense heritability. For calculation of narrow sense heritabity, crosses have to be made in
definite fashion. It plays an important role in the selection process in plant brecding. The main
Teatures of narrow sense heritability are presented below.
1. Application: It is useful in both animal breeding and plant breeding. It is useful ia
sclection of elite genotypes from segregating populations,
2. Estimation: lt is estimated from additive genctic varance,
S. Formula Used : It is calculated with the help of tollowing formula
Heritability (ns)=1/2 D/VP, where, D is additive genetic variance and vP 1s phenotypie
vanance.
Since narrow sense heritability is estimated from additive (heritable) genetic variance,
plays important role in the seletion of elite genotypes from segregating populations, whercas
broad sense heritability estimates arc useful in selccton ot chtc genotypess trom homozygoun
materials. Main differences between broad sense heritability and narrow sense heritability a
in Table 9.1.
prescnted
TABLE 9.1
Comparison of Broad senseand Narrow sense Heritability
S.NParticulars Broad sense Heritability Narrow &ense Heritability
Vaniance used tor estimation Total genetic variance Additive variance
Populations used parental
Both
segregating Specific populations
and
Useful in selection of elte Homozygous lines Segregating material
ypes Trom
Animal breoding
4Application in Both animal and plant Dree
ESTIMATION OF HERITABILITY
As stated above, heritability is estimated in two ways, viz. broad sense and narrow sense. N thod
of estimation of heritatbility are presented as folows.
1.In Broad Sense Broad sense
heritability from simple replicated trials and
cstimatcd as follows. gene
mcans 1s
Genetie Adrac
and
iy
Simple ple Trials :
Heritablity e)-VY e9
Frem
phenotypc warai,fepetvdy m 97 gy
eneration mean analyis zHertubiley e)V2VPPn , 97
V E 2 are variance of Fl and P2, tepectiney
Sense : Naow sene tersatiliey fass dild s n d gooon s o
follows
ron
Disllel AnalysisTbe ollowing ormuls js for sting
beritahility from diailiel cnoes.
Fl
Henuability (as)=|1/2D +2 HI-12 H212 FASAD2H-MRDNR
etalen and Murray (9)poped te foliowing fomda Sor sneing
beritability trom F geeraaot a dialied c o
ot
Herntahility (nsj= 14DA1/4D -1/i6 HI- F-E
Mes Anadysis : fNamow sctue
m From Generation teiliny ie sined oalows
As per Warmer (1923. Hertatiny = 2 j 4 , wec.D 92RE SOA
and phenotYpuc waranoes nespecy, Oniy D s s u so cliine s n s
heritability. because the phentypic atiatce sf P2 tus soy 9Dwu additos cmgn
As per Mather (3909), Beritabiliny= D D HB 3 wm d E
additive, aot-a0one
er .
Scales
Scarie (1965) has suggesied minitnum vaiues of samow eatiiny s
alinect seiectios to be mone efficien that diec aieinae Sor yoeit. Te wale d
ad broad sense heritaility are peesetted it Tatic 922
TABLE 92
Scales for Bruad sense atd a r sse heritathlley
Narrpw sense (Searie. 1965) So
Beiow 6P
41 1o 50 sleiu
Above 50 g
ACTORS INFLUENCING HERITABILITY
i e s of heritatility ae infiucaced by vati facte. ge gseic mun
G
si, ) samgiling ssethod, 14)zondu f cngetimet mtid arf caicatition.a
L
(Satonde, 1979, Febe. 197 hec fao efiy daca
Genetie Materiad The magnitude af hernnin
e p i c variance presen in the hding pupuliain fr t
y p i c varianoc ighor the heriatieg The deta f dg p
88
Esentials of Plant
closely related. Bri
are
expected to express more genetic vanance than derivatives of
degree of inbreeding also influences the genctic vanance. The
increase in the rate of genetic variance i
inbreeding. Thus, hentability estimates are
after inbreeding. higher in F4 or pe
2. Sample Size :
Henitability estimates are influenced
estimates are based. If the estimates are based on the entire by the sample size on
variance of a population, but evaluation of entire population, it will give tm
sample will give estimates near to the POpulatton 15 not practically possibie nds
population mean, while small sample
information. may give m
3. Sampling Method: There
are two main
masieading
biased sampling. The random sampling methods, vEz, random
vaniance and hence sampling method generally provides truc sampling e
henitability. The biased sampling on the estimates
other hand will
of gme
representative estimates of genetic variance and not give t
of
genotypes is evaluated to thereby hentability. when a
the ratio compute the ratio of
genotypic vanance to the
non-random sampe
is not called phenotypic varianc
heritability. This is called
repeatability.
4. Conduct of
or error
Experiment: Heritability estimates are
variance, because eror variance influenced by the
estimation of part is a
part of of environmental em
heritability. The degree of uniformity in the testphenotypIC vanance used in te
experimental error. Any precaution that a breeder can environment will
influence te
the take to reduce
improve heritability of a character. The error vanance can be expenimental erTor (VE) wil
plot size or number of reduced by either
estimates of replications. Moreover, multilocation testing will
increasing te
henitability than testing at one location
only. provide more reliatie
5.Method of Calculation :
Heritability is estimated by several methods. The
heritability obtained by vanous methods
heritability are always higher than narrowwill vary to some extent. TThe estimaies
sense
hentability.
estimates of broad sctt
6. Effect of
Linkage: Heritability estimates are
linkage. Linkage influences the heritability considerably influenced by the presence
estimates by causing an
the estimates of additive and
dominance genetic vanance. upward or downward bua
linkage (AB/ab) causes an upward bias in the High frequency of coupling
variances (Hallauer and Miranda, estimates of additive and
1981). An excess dominance
upward bias in dominance vanance and downward biasof repulsion phase linkage (Ab/aB)
can be reduced in additive variance.
by random mating of
population. The number of Linkage discqu
required tor breaking the
linkage depends on the closeness of the generations of interm
linkage (hanson, 1999
Advantages
1. Estimates of henitability are useful
in predicting the transmission of
to their otfspring. characters from p
2. Estimates of heritability
are free from genetic assumptions
3. Heritability estimates are based on
empirical results.
4. Broad sense heritability can be
calculated from both parental as well as segres
populations5.
5. Heritability estimates are useful in selection of elite
as well as from segregating populations genotypes from homozygous t
G e n e t i c
vance
Advan
an
hilioy
eritability gives an 1dca about additive genetic variance
6 N a r r o wS e h s e h e r i t a b i l i t
n i t a t i o n s
jn 1.
ates
E s t i m a t
of
Tooust
On second order statistics (variances) and, therefore, are
heritabilty are buseureliable.
and
not
statisticauuy vei
ance and its componcnts (additive, dominance and epistasis), as compared to
he v
2ns are also inaccurately estimated.
of variance sometimes turn out to be
Componctnts
negative, resulting in negative estimates
of heritabiny.
Heitability is a
t0
property of a specific population in a specific experiment. Thus heritability
Eenetc material,
is specnc
estimates of heritability are influenced by several factors such as type of genetic matenal,
The e
ample size,
sampling method, conduct of experiment, method of calculation and linkage ettecs
1979: Fehr, 1987). The random sampling method provides true estimates genetnc
Summonds, (based
hence of hentability, when hentability is estimated from non-random sample
ance and
of notypes, it 1s caled repeatability.
ample
COHERITABILITY
in
Co-heritability 1s theC Tato ot
Eenotypic covanance to the phenotypic covariance expressed
ot co-
t1s measure simultancous innentance or twO characters. Hence estimates
percentage. o 15
heritability are useful 1n simulaneous improvement of two characters. The co-heritability
estimated with heip ot Tollowing tormula.
Co-heritability (x, X,)= GCOV X , c o v X,X 100, where GCOV and PCOV are genotypic
nd phenotypic co-varlances of characters X, and A,. respectivel
values
High of
co-heritability that increase in
estimates suggest one polygenic trait will lead
simultaneous increase in another co-heritable character.
TABLE 9.3
Comparison of Heritability and Co-heritability
SN. Particulars Heritab Co-neritability
1. Trait Involved One wo
Degree of statistics invoives Second order Second order
Single trait wotraits
Useful in improverment of
5.
Types Two only
Accuracy ot results Mediumn
Medium
GENETIC ADVANCE
he n
wement in mean genotypic value of selected plants over the parental population is
tnetic ad netic
enetic advan advance. It is the measure of genetic gain under selection he success of
d se c e under selection depends on three main factors, VI. genctuc vartability, heritability
don
intensity. These are explained as
TO ows
Essentia of Plan
1. Variability
The magnitude of genetic
: variability present in the base Br
poant bearing on the genetic advance. The genetic advance is expected t
Population having moe genetic divenity be hip
2. Heritability: The heritability of character under
consideration affects geneti
Thegenetic advance is expected to be high in those character having high values of t
3. Selection Intensity Selection Intensity refers to the proportion of herit
seiected from a genetic plants
population for study. High selection intensity also gives better or t
low selection
intensity. Genetic advance is estimated with the help of
following formeds
et al, 1955).
mla 0ale
Genctic Advance (GS)= (K) (H)
(SDP) or GS= VgVP 1/2,
under selection, K is selection
differential and SD P is the where, GS is genetic
population and H is the heritability of the character under phenotypic standard deviatioe of
generally would give higher estimates of GS than seliection. Broad sense herit
advance are rated as
low (<l0%), medium practically realized. The values of g
(10-205) and ( high 205).
Scales
The minimum values of genetic advance
than direct selection for
yield have been
necessary for indirect
selection to be more effioes
arbitrarily suggested as given in 1able
TABLE 94 94.
Arbitrary Scales for Genetic Advance
SN. Values of Genetic Advance
Less than 10 (percent) Hate or Scale
10 to 20 Low
Above 20 Medium
High
GENETIC GAIN
The difference between the
mcan phenotypic value of the
population is known as genetic gain. ft ts denoted by R. Itprogeny of selected plants and the
bo
R-XP-X0, where XP and X0
and base population, respectively are the mean phenotypic
is calculated as follows
values of progeny of selecsol
Selection Differential
The diffcrence
population is termedbetwcen
as
mean
genoy pic
selcctbon
valuc
of
progeny of
selection Selcction lnichsily differential. It is denoted by K. selected plants o
It is the measure
renetic population 1or study retcn to the of the
selected individuals 1he value of K proportion of plants or
generally CAceed the meanvanies with the intensity families scleci
s calculated as follows of parental of selection. The
K-XSX), where AS and A) ae the mean population The selection dit
population, respectively. phenotypic values of
selected plants an parena
Genenic Advance
and
ay
ance from mixtures ot
lines
ecnetetimates and pure or cloncs should be
e hentabilityye trom
secgregating populations using narrowcalculatod using bec
sense heritbsly
AdVantagess
Estimates of a enetic advance are based on
ctnpinical results and are free
assunpto
fromg
is reliable measure ot
genetic inprovement under selection for
it
polygee
advance advance can be
can be eestimatcd trom
both parental as well
Genetic as
segregating popa
genetic advance nelp in
4
mates o of
understanding the
type of gene action involved in the
ion of various polygenic chiaracters,
High values of genetic advance arc i n a e
additive gene acuOn and eOw vaues are
indicative of non-additive gene
action.
walso helps in deciding a Dreeding procedure for the genetic improvement of
van
polygenic traits by determining the gene action.
Disadvantages
1Estimates of genectic advance are based on second order statistics (variances) and. therefore
are not statisticaliy very robust.
2 Estimates of genetic advance are specific to the breeding material. Hence. estimates vary
according to breeding material
INTERPRETATION OF RESULTS
terpretation of the
Cssed as ollows
results of heritability. genetic advance and combined results of both are
Heritability
value of heritability in broud sense is
the high, it indicates that high value is due to
favorable effects of environment. Broad sense heritability is calculated fron total genetic
varance which include both fixable (additive) and non-fixable (dominance and epistatic)
varzances. Hence, selection for characters showing high values of heritability may not be
etfective
h e value. of broad sense heritability ts low. it reveals that the character is highly
cnced by environmental effects and the genetic improvement through selection would
iCult due
masking effects of environment of on the genotypic effects.
if tability estimates are high in narrow sense, it means that the character is largely
EOveriedby additive genes and selection for improvement of such character would be
rding.
If
non-
heritabil estimates are low in narrow sense, it shows
that there is
preponderance of
e gene action and heterosis breeding may De useiu
92 Essentials of Plant
TABLE 9.5
Bred
Efectofselectionwith high and low values of heritabilityand geneticaadvance
S.N. Estimates Values Expected Gene effects
Selection
Heritability (bs) High Additive& non-additive
2 Heritability (bs) LOw Non-additive Ineffectve
nefecie
Heritabilily Hign Additive Efective
Heritability (ns) LOw n-additive neflectve
Genetic Advance gn Additive fective
Genetic Advance n-additive
eneci
B. Genetic Advance
1. If the value of genetic advance is high, it shows that the character is governed by additive
genes and selection would be rewarding for genetic improvement of such trait
2.
the value
Tfaddituve of genetic advance is low, it indicates that the character is govemed oe
gene action and heterosis breeding may be useful.
by
C. Heritability and Genetic Advance Combined
High heritability accompanied with high genetic advance: It indicates
likely the heritability is due to additive gene etfects and selection may be ettectuve
that mo
2. High heritability accompanied with low genetic advance It is
indicative
: of noe
The high is being exhibited due to favorable influcnce of
additive action.
gene
environment rather than
heritability
and selection tor
genotype uch traits
may not be rewarding
3. Low accompanied with high genetic advance: It reveals that the characier
heritability
1s governed by additive gene etlects. The low heritability is being exhibited due to bigh
environmental effects. Selection may be ettectuve in such cases
4. Low heritability accompanied with low genetic advance : It indicates that the characte
is highly influenced by environmental effects and selection would be ineffective.
Ouestiens
