DBDl.CATION

To my rn'Ífer Yulizar Sutedjo, and mY

chíldren, Anita, Linda, A¿rdy and Chusnul, for

their moral support, unclerstanding and

enc.ouragement.

J-

STATEMENT

reference Ís uade Ín the text'

T-?.

ACKNOI,JLEDGEMENTS

I wl-sh t'o express my síncere gratiÈude to my SuPervisor,

Dr. R. Knight, for hís advíce, guÍdance and crÍtícism throughout

thís study.

I woul-d also like Ëo thank Prof. C.M. Donald, Dr. R.J. Rathjen

and Dr. K.W. Sheperd for valuable advÍce relatlng to the study'

Mr. T. Hancock and other staff in the Biometry section gave'

considerable helP with comPuting

Many rnernbers of the Agronomy DeparËment at the I'Iaite.Instítute

assisted the author during the conduct of the consecuËive field

Èrials in 1975 and 1976.

Fínally, I an graÈeful to the Conrmonwealth Government of, Australia

for flnancíal support through The Australian Asían Universíty Co-

operaÈion Scheme and to the Government of the Republic of Indonesia

c.q. departmenÈ P & K for offering the opportuníty to study'

¿¿t

SIJ}OÍARY.

1. Fíeld experiments rntere- conducted in 1975 and 1976 to Ínvestigate

the efficíency of mass selection ín ldimrnera l:ye.gl:ass ("LoJium

rigidum.Gaud). Populations of 3,136 planÈs in 1975 and 3,920

plants in 1976 ì/ere gïohrn "t a àeosity to símulaÈe swards in

areas of 8.75 m x 8.75 rn and 8.75 m x 9"50 m respe-ctiveIy. To

take account of local variation within these areas, they were sub-

dívide-d into plots of .875 m x .875 rn. Each population was truncated

by selecting the three- highest yielding plants out of the 49 plants

in each plot. Yield r'ras neasured as the total dry matte.r produced

at three harvests taken approximately 30 days apart. The selected

plants rrere moved to a glasshouse before anthesis to facilítate

intercrossing among them. In 1975 , lrg2 plants were selected and

moved to the glasshouse.

2. T\^renty offspring from each of these sel.ected plants were grown in

a field trial in Èhe second year (1976), together v¡ith unselected

material that served as a control. This trial revealed that the

selected material had a 13.8 per cent yíeld advantage on average'

over Èhe controls. This íncrease hacl been achíeved by one cycle of

mass selection.

3. Because the distribution of the populations \,üas skewed, the effect

of mass selection was analysed by using the percentage of the

plants below and above the mean and also below and above the mode

of the population. The results of these analyses showed that the

population derj-ved from selec¡ed materíal was less skewed and

more normal than the unselected population.

4. A procedure for assessí-ng conbining ability tlas suggested and

studied. IË was based on the number of offspring of each se-lected

plant which in turn would have been selecte.cl for further breeding.

5

l.v

If the off sprirrg of the selected. planÈ appearecl at a high

frequency in the truncation of the following mass selection,

the selected plant had a hígh general combining abilíty. On

the other hand, if they occurred at å'low frequency' it r¡as

suggested the selected plants had a hÍgh specífic combÍning

abil-ity. Reason ín support of this suggestion are presented.

Plants \^Iíth å htgh general combining abílity were regarded as

potential constituents of a synthetic variety. Rese::ve seeds of

such plants could be compounded and released as a synthetíc

varÍety.

An attempt to produce selfed seed t/as unsuccessful. This could

not be attributed conclusively to self-Íncompatibility. It 4ay

have been due to unfavourable environmental condition wlthin

the pollination bags.

Alternative procedures to mass selection, including the combínÍng

abillty assessment índicated in this study are presented and

discussed.

6.

v

CONTENTS

STATEMENT

ACKNOI^II,EDGEI"IENT S

SUMI"IARY

I,IST OF T'IGURESLIST OF TABLES

INTRODUCTION

1. Mass selection2. Theoretícal view of mass sel-ection3. The mass pedígree method ín corn4. Selection in forage crops5. Al-ternatíve breeding procedures in w'

ryegrass (e.g. synthe-tic varieties)6. Theoretical view of synthetic varíeÈies7. Micro-envíronmental vari-ation and the

selection of genotYPes

EXPERIMENTAL DETAILS

SiteClÍmateSeed sourcePlanting preparaÈionPlant spacingFer:tilizingHarves È

YieldField layoutThe selection sequence

RESULTS

lsgg.

I¿J

-t¿Jvlt

v¿r.¿

I

45I9

1011

t2

1234567

89

l0

151515151B1B181B

1919

23

31

33

1. Yíeld results2. Affect of selection on the skew of Èhe

population3. Environmental heterogeneity withín the

experimental areas4. The distributions of yield in populations

of mass selected Plants5. Progeny test in w. ryegrass6. Self-pollinated seed

DISCUSSION

1. Mass selection in w. ryegrass2. Plant assessments3. Synthetic varie.ty in w. ryegrass4. Alternatí've mass selection procedure in

I¡7. TyegrAsS .

5. Purpose and expectation from the thírdyear testing

363639

4B

5051

52

52

YT

CON1ìENTf'l ( con tirrrrerl)

REFERENC]JS

APPENDIX ].

APPENDIX 2

APPENDIX 3

Yielcl in 1975 (total of 3 l-rarvests)

Yierlrl in L916 (total of 3 har:vests)

Yiel d of the 20 of f spríng (srrm of 3

harvests) I976

3-ag-".

5l

63

-/r

B1

.t

\'l

ll

var

Figure

LIST OF FIGURXS

Mean monthly raínfall in mm

Layout of the experinent in 1975

Layout of the experiment in 1976

4a, b & c Distribution of yleld for the lst, 2ndand 3rd harvest ' L975

5a, b & c DísËríbution of yield for the 1st, 2¡dand 3rd harvest, L976

Page

20

2l

24

26

27

37

38

I

2

3

16

25

34

6a&b

7a&b

8a&b

9a&b

10a&b

DistributÍons of Èotâl yield for Ëhe

harvest in 1975 and L976

Distríbution of yields for the origlnaland selected population 1976

M,icroenvironmental varíatíon based on themean ¡zields of the 49 plants ín eachsub*plot in 1975 and l-976

Distributlon of selected plants based onsÍmple and subdlvj-ded plot truncation inL975

Dístributíon of selected plants based onsimple and subdivided plot truncation inL976

I

I

ii:l

jl

f,i

,ì

I

VJ.JI

Table-

4a

4b

6b

10a

10b

2

1

2

3

I,IST OF TA]]I,b]S

A description of the characteristics ofsoïne cïoss-poI1inaÈed ctops includingl^7. r}egrasS

Clím¿rLologíca1 val-ues during the yearsof Èhe experínents I975-L976.

The date of transplanting and harvesting'

The mean of the total yields for threecutting 8/Plant drY matter.

Total yíeld from three cuttings in kg/hadry maiter (plant density 640,000/ha) '

The heritabilítY of the Yield of w'ryegrass in Lhe lsË mass selectíon

The percentage of the plants below and

above the mean of the PoPulatíon

The percentage of the planËs below, above

and at the mode of the PoPulation

The percentage of the planl-s below themöde and the mean, and the skewness ofthe population

The values of mode, meán and skevmess ofthe population

The frequency with which a selected plantrsoffspring occurs among the highest yíeldingplants in the next generation. (Max'

possible score 20).

A rankíng of selected plants on a basis ofËheir yield together wj-th the frequencyof reselection of offspring and the 20

offspring mean yield

A rankíng of the sele.cted plants based on

frequency of ::eselectíon of offspríngtogether wiÈh theír yield and 20 offspringmean yield

A ranktng of selected plants on a basis ofthe 20 offsprj-ng rnean yield together wiEhtheir yield an<l frequency of reselection

Page

40

4I

43

5

6a

7

L7

1B

29

29

31

32

32

32

338

9

I

I

10c

4s

1

WAITE INS'i:li.i f¡:

LIBItAiìI'INl]RODUCTTON

This thesis ís concerned v¿ith a stucly of mass selection in

I¡iimmera ryegrass (LoJium rigidttm.Gaud) á aiptoia, annual cross-

pollinated grass species, It also deals with a possible alËe.rnatíve

breecling pr:oceclure to mass select-ion (e.g. a form of progeny Eesting) '

In any plant breecling pïogranììme, the selection of desirable

genotypes depends on the char:acteristics of the species with r¿hích the

breeder is concerned. Each species has a characteristíc lífe cycle,

flowering period, reproductive system, pollinaÈing mechanism and other

attributes which affect the choice of techniques. The choice is

further affected by consideratíons of time, sPace and l-abour. Breeding

procedures may be considered as steps connlencing with

(1) building up the population to be selected'

(2) selection itself' and

(3) maintaining and nultiplying the selected plantsfor comhercial ProductÍon.

For the first ar-rd the thÍrd steps most breeders are in agreement;

they attempt to create populati"ons having the desired combinat-ion

and frequencies of genes and to economise in the nr.rmber of generatiorrs

required when multiplyí.ng the selected plants to be released as ne\^I

cultivars. But for Ëhe second step' breeders differ in their ideas

of the most efficient procedures and Èhese depend to sorne extent on

the characteristics of the cïop. To place in conte-xt the lnass selection

considered in this study a description of the characteristics of some

cross-pollinaterl cl:ops including I^l . l:yegrass is given in Table 1.

In corn self-pollination by hand is simply affec-te<l and this

facilitates the prodrrction of hybríd and synthetic varieties. However,

its yíeld evaluation (the grain yíeld) can be rnade only after floweri-ng

TABLE 1 SOME FEATTIRES OF CROSS-POLLINATED PLANT

Plant Species Life CYcle

Annual

tr'loweringI'requencY

Once iìr aliie cycie

PollinatingMechanism

Self-compaÈible

Reproduction System andics EfficiencY

Seed only Efficient

Yield EvaluaÈion

After flowering(Grain)Corn

(Zea nags.L)

Sugar Cane(Saccharumcfficinarum L)

Coconut Palm Tree(Cocos nucifera. L)

Cassava(Manihot utifisima.L)

Once in alífe cycle

Self-incompatible

Biennial

Perennial

Annual

Once in alife cycle

Self-incompatible

Seed

Mainly vegeta-tively propagated

Seed only

Seed

Mainly vegeta-tively propa-gated

InefficientEfficient

Efficient

InefficientEfficienË

Before flowering(Sugar content)

After flowering(Copra produc-tion)

Bef ore f 'l owering(Tuber produc-tion)

More thanonce in alife cycle

Self-cornpatib 1e

Perennial

Annual

More thanonce in alife cycle

Mainly self-incompatible

Seed

Could be vegeta- 'tively propagaÈedfor seed produc-tion

Seed

Could be vegeta-tively propagated

Efficient Before flowering(Forage produc-tion)

Before fiowering(Forage produc-tion)

Cocksfoot( Ðactq 7isglonera.ta. L)

tr{immeia Ryegrass(Loiium rigidumGaud)

It

Once in alife cycle

Self-incompatible

EfficientComrnerciallyInefficient

NJ

3

wlrereas Ín W, ryegrass, sugar cane and the perennial grasses, yield can

be evaluated before floweríng. !üitir the latter: cr.ops there is an

opporl-unity to select and cross among the selected plants to produCe

seeds in every generation before the 1ífe cycle has been completed.

I,liÈh corn, breeders would be able to select in each genelation only

if there Trere a good correlation beÈween yield and some characters

before flowering. No such correlaÈion Ís known. In sugaT cane a

desirable genoËype is use<1 ín a crop after vegetatíve propagation.

The portion propagated develops ínto a nevT plant with a rener,¡ed life

cycle \nlhereas ín W. ryegrass, rice and wheat, the porÈíon of vegetative

organ ís a tiller, and the life cycJ-e is not renewed. In many perennial

grasses such' as DactgJis gTorerata L, ít ís easy to maintaín the

parents over several years and to make repeated crosses.

To summarize, w. ryegrass is an annual ín whÍch a yíeld assessment

can be made before anthesis, so that a cycle of selection may be made

each year. It carÌnot readily be vegetatively proPagated and parental

material cannot be maintained from one year to another. It ís riot knolvn

whether self-polJ-inaÈíon would produce suffícient seed to be useful in

breeding buÈ this seems unlikely as the species is cross-pollinated.

Furthermore, íts flo\,Jers are hermaphrodi.te unlike corn and the control

of pollination used to produce hybrids ín corn is not a feasíble

alÈernative Ín vr. ryegrass. The progeny tesËs used in corn could be

difficult to undertake and mass selection is an obvious breeding pro-

cedure to study in w. ryegrâss.

4

LITERA]'URB IìEVIEI^'

Many selection p::ocedures for cross-pollinated crops are base-d on

genetic concepts developed by corn breecle.rs in the U.S.A. These con-

cepts ínclude combínirrg ability, hybrid vigour, inbreeding depression

and herítability.

It is informative \nlhen attenìpting to improve the yield of other

cross-polU-nated cïops to consider the relevance of corn breeding

procedures.

1. Mass selectíon

'Ihe objective in mass selection ís to increase the proportÍon

of superior genotypes Ín a population by undertaking selecÈion on

a basis of the phenotypíc value of planÈs in Èhe population. Its

efficíency depends primarily on gene frequency and heritability.

Generally, híghly heritable characters respon<l well to mass selection'

In corn, it was possible to develop varieties differing i.n grain

colour, plant heíght, síze of ear, placenent of ea¡ on the stalk, date

of maturíty and percentage of oil and protein content (Sprague, 1955;

Allard, 1960). These are all highly heritable characters' Characters

with a 1or¿ herítabil-íty such as yield gave variable results. For

yield, a 38 per cent increase has been obtalned from Èhirteen genera-

tions of mass selection of the llays Golden variety (Gardner, 1969)

and in Mexico a 33 per cent increase I¡Ias obtained from three cycles of

mass selection Ín a tropical corn variety (Johnson, 1963). However,

Hallauer and Sears (1969) ve-re unabl-e to show significant improvement

of yíeld in either "Krug" or rrlowa ldeal" flom slx cycles of mass

selectíon.

In other crops also, such as sugaï beet and alfa1fa, there is

5

evícle1ce. that irnprovement fron .long-cont'i nued mass selectlon rvlll

improve. highlv heriÈable char:acters (A11ard, 1960) " On the other

hand mass selecti-on has been less efficient in modífying characters

such as yie1d. This characler is gove::¡ed by many genes and its

expression is influenced by the envíronment. Single plants cannot

often be judged acculately for their breeding value on a basis of

their appeararÌce.

2. Theoretical víew of mass selection

fn mass selection, clesi-rab1e individuals are chosen, and their

seed composited withouË a progeny test to produce the following genera-

tíon. The selection is base-d on the maternal plant and ther:e is no

control over pollinaËion (Sprague, 1955)

Mass selection for a single gene ín corn e.g. a gene for kernel

colour is sirnply undertaken by selecting kerne-l-s v¡ith that colour.

Each kernel contains a gene for that colour. But selection of the many

genes that control plant height, cannot be undertaken mere.ly by taking

kernels from plants with the desired height, unless the plant ís a

homozygous ínbred for that character and environmeni has not been t,he

maín factor determinj-ng height. In the stage of gamete formaÈion the

probability of the height genes occurring ín the fernale. gamete will

depend on heterozygosity, multíple allelísm ancl polygenic inherítance.

In addition with mass selection, the corn plants are cut-pcllinated

and the male parerrt will not have been selected for height. Therefore

the chances of selecting erfficiently t-he genes for height in corn ar:e

remote. It could be done only by selfí¡g or by transplantíng the

selectecl plants before anthesis to an isolation block' Such a procedure

would be very laborious. Even rnore difficult is the selection and

6

utj-lisation of high yielcling genotypes. Yiel-d calìnot be assessed

before anthesis. Iurthermore' not all the kernel-s which are derive-d

from a high yielding corn-cob will contain genes for high yie1d.

As yield is the end product of the physiological developmenL of

a plant in íts environment, any gene involved in cle-velopment must be

a yi-eld ge,ne (Shebeskí L967). Shebeski concluded that hundreds or

thousands of genes musË be involved in the yield of crop plants.

The difficulty and ineffícíency of mass selectíon for ylelrl in

corn results from Èhree main causes:

(i) an inabilíty to ídentify superior genotypesfrom the phenotypic appearance of singleP lants

(íí)

(iíi)

uncontrolled pollínation so that seleCtedplants are pol1ínated by both superíor andinferior planÈs, and

more rarely a too high selecÈion intensityresulting ín a reduced population size andinbreeding depression (Lonnquist, L949 ;Allard, 1960).

Mass selectíon for forage yield in w. ryegrass may be different'

Selection can be carried out before anthesis which enables the inter-

pollination of both female and male selected planÈs. Therefore the

expectation for a change in gene frequency accompani-ed by an improve-

ment in the mean of the population is greater.

The efficiency of mass selection fs influenced by the type of

gene action that ís determining the characters under selection. If

yíeld is determined by gene actíon r¡hich is additive, completely

dominant, cornplementary or conglomeraÈe (Cockerham, 1956) nass selection

in w. ryegrass should íncrease the desirable gene. freqrrency. But íf

yield j-s determined by the heterozygor-rs condition and overdomin.anc.e

the¡ mass selection r^¡ould select heterozygotes and the.re would be no

7

change -i-n gene frequency in the next generatíon. Inlhat evídence is

there for thinking that yíe1d is deteruined by one type of gene actíon

or another?

f.n a recent ïevíe\¿ü of gene action and heterosis (Sínha and l(hanna,

J:g75), slrpport was pïesented for an hypothesís pTesented many years

earlier (Keeble and Pellew, 1910; tr{ílliam, 1959). According to thí-s

hypothe.sis, heËerosis is the result of the multíplicative natule of the

components of yíeld. If heterosis for yield is analysed aË the- 1evel

of its componenÈs, the results for the components show a simple inheri-

tance pattern. Each component of the hybrid resembles eíther parent

or is intermediate bet\^reen its païents. However, because the components

are multj.plícatíve the yield of a hybrid rnay exceed that of its parents.

If this hypothesis is coïrect Ëhe inheritance of yield conforms with

the donínance hypothesls (snutt and EasE, 1908; Brewl¡aker, L964;

Strickberger, L976>.

According to this theory, we should exPect pure heterotic lines

to occur which are hontozygous for dominant alleles at all locj-.

However, although such lines have been sought oveï a long period of

time they have never been obËaíned (StrickbergeT, 7976). The reason

for this failure may be the large number of genes affecting yield and'

Itnkage of ïecessive genes to the favourable dominant genes. The

absenc.e of homozygous heteroLic línes therefore, does not necessarily

preclude Ëhe dominance hypothesis.

Based on the evidence given by Sínha & Khanna, a hybrid would

exhíbit heterosis r¡hen the components of yield are inherited as linked

domÍnalce-recessive factors ín one pareDt and these are compl-ementary

to factors in the other ParenL.

I.lowever, to confirm a dominance hypothesj-s in a polygrric character

B

such as yíe1d, it rvoul.d be necessary to have- extremely large populal-íons

for the-re to be the poss:t-bítity of the genotype that j-s horoozygous

for dorninant al,leles at all loci to aPpeaï in the F, PoPulation'

An approach to this problem of large popul,ations ís Ëo use a iype

of popul-ation ímprovement which will effec.tively shift the frequency

of desirable- genes tov¡ards homoz¡'gosity (Genter, L967) '

In another study, hybrid vj-gour was related to genotype-environment

interactions (Knight, 1973). The possíble relaËion between ':he Tesporìse

of two parents and their F, when groÌ¡/n ín various enví::onments \'{aS

presented. Where a hybrid had a response exactly intermediate

beÈween its parents, metric values for the hybrid \nrere not intermediate

buË varied with the environment, exhibiting di.fferent degrees of

dominance including overdomínance (hybrid vígour) ' Thís interp!-eta-

tion is to some extent in agreement r,¡iÈh the suggestíon of shinha

& Khanna in that it should be possible to obtain ge-notypes as good

as the F, frorn selfing the hYbríd.

The genetic basis of heterosis remaíns unresolved but the con-

sistent ínc,rease in yield through mass selection 1n corn (Gardner,

1961), supports Èhe idea that selection has led to an íncreasÍng

desirable gene freq\rency in the population and that .heterozygousity

and overdominance aïe not the main or only forms of gene action

determining yie1d.

3. The mass oedisree method in corn

This method involves Ëhe addition of a pedigree plogeny test

to mass sel-ection and therefore has been called the "Mass-Pedigree

Method" (Harrington, Ig37). After mass selection the selected plauts

are open-pollinated ancl a progeny test carried out with one family

9

in each plot (Macaulay, 1928). I^lj-th this proce<lu¡:c Ëhere is an

assurnption that a lorv le',zel of inbrecding takes place because the

pl.ots are païtia11y isolated acco::cling to family ancl also because

soue of the kernels on each ear aïe. naÈura1.ly self-po1-1Ínated" The

value of the inbreecling ancl sib-pollination is assumed to lie in the

slower approach to homozygosity than would be tl're case with totally

controlled inbreeding. With thís proceclure there are greateT oppor-

tuníties for gene re-conbir¡ation ancl provision of an oppoTtuniLy for

elímination of some of the less adapted individuals thror-rgh natural

selecËion. This procedure could be applied to w' ryegrass'

Other modífications to these breeding procedures are summarized

by Andrus (1963). He stated that the best features of pure line selec-

tÍon combined with several generaÈions of mass selection with natural

or enforced sib-crossing early in the breeding cycle shoul-d be effec-

tive in improvíng Yield.

4. Selection in forage croÈq

In a review of selection methods in the breeding of cross-

fertílíze<1 pasture species l,atter (1964) staÈed that the characters

of economic importance ín a pasture species are predominantly quantl-

tatíve. To evaluate quantiÈative charactels in a breeding progranme'

a scheme of progeny tersting was needed. In some cross-pollinated

crops there are no dífficulties because it i.s possible to maintain

païenËs and progeny. But as previously mentíoned a cortventíonal

progeny test for w. ryegrass cânnot ieaclily be undertaken'

In a recent study of breeding for forage yield, an improvement vtas

obtained using a modified form of mass se-lection called "Rec.urrent

restricted phenotypi.c select:j-ort" or "RRI'S" (Burton, 'L974). The- procedure

contained five features designed to Írrcrease íts efficiency:

10

(1) A space-planted population \ùas dividecl intoplots each of 25 pJ-arrts. This was done toreduce the effect of soil heterogeneity on theselection of plant.s" The fíve- highest yieldiugplants among the 25 plants ín.a plot wereselected as parents for the ¡rext generation.

(2) 1'he selected phe.notypes were j-solated beforef1-owering, enabling paternal as wel-t as maternalselection.

(3) Inter-mating was facj-litated by mingling theflowerÍng culms from each selected phenotype.

(4) By using two heads only from each selected pheno-type the autbor avoicled unequal- representationof parents in the next cyc1e.

(5) And by choosing gerniplasm wi-th a high degre-e ofself-incornpatability he v¡as able to r.educe thelikelihood of selfing.

BurLonfs study was with Pensacola bahiagras (Pasþalum notatum vat.

saure "Parodi") a perennial which can be propagated vegeÈatively.

Using a selectíon íntensity of 20 per cent in four cycles of RRPS,

there \¡/as a forage yield improvement of 16 to 19 per cent over the

commercíal check varieLy.

To some extent this procedure could be applied to w. Tyegrass'

buÈ ít needs some modificaÈion because of the dtfficulties of veget,ative

propagation.

5. Alternatíve breedin rocedures in w. e rASS S ntheti cvariet-ies

Synthe-tíc varietíes are developed ín corn by inter-c::ossing

several genotypes. Ìor other cross-pollinated crops which can be

propagated 'vegetatively, selected clones could be useci.

Gallaís et a1. (1970) defíned a synthetíc variety as an artificial

population ín which the successive generations result froin natural

crossing of clones or farnilíes chose.n as much for theír physiological-

characteristics as for general and specific cornbínÍng ability.

e.

11

Íurthermore, he stated Lha.t if the c,onstl-tuenLs aÏe famílies, thcric

may be inbre-<i 1ines, single or dorrble cÏosses ol: even ecol-ypes of

clÍfferent orígín

It will be- noted that tl-ris def ínj tion of a synthetic vari-ety

does not r:equi-re that the línes or clones in the synthetic have bet'-n

tested for conbj-ning ability. However.,'Allarc1 (1960) defj-ned a

synLheÈí.cvar:ietyasoneÈhatwassynthesizedfromgenotypesr¿lrich

have been tested for combii-ring ability'

6. Theoretí cal view of svnthe.tic varieties

SyntlreÈicvar:ietiesincorrrwerefirstsuggestedbyHayesan<1

Garber (1919). In general these early synthetíc varieties had yíe1c1s

1ìo greater than open-pollinatecl varie-ties. But later when the con-

sËituenË genotypes in the synthetic variety were chosen on the basis

of the.ir combining ability, the yíelds of synthetic variety increase<1

ancl were superior to the open-poll:Lnated varieties (Hayes et a1' , L944) '

Twomethodshavebeensuggestedforj-ncreasingtheyieldof

synthetics produced from several inbred lines. These are by increasing

the yield of the inbred itself and by increasing the F.'s mean yields

(Kir:nan, sprague 7945). Tn corn this can be accomplished by fixing

the desírable gene or gene combination by inbre.eding, selecting among

the inbreds, followed by progenty testing rvÍth a view to selecting

the inbreds that provide maximum heterosis after synthe'sis'

Inaddition,t'henumberofthegenotypesinthesyntheticshould

be considered. Gallaís (1970) ga'\/e a formula for the vigor of a

synthetic developed for a diploid species. The spec.ies ís assumed to

have incomplete self-incompatib'i-l-ity, random rtating occ'urs, atrcl there

ís no competition betwe.en planLs.

Inpracticeitispossib].etod.istirrguishtwogroupsof

l2

cross-poll-j-nated crops; those whe::e the comuercial production of

F. hybricl seed for farmer:s ís feasible, ancl those in r¡hích further-l.

multiplication of seed is necessary. \nI .' ryegrass bel,ongs t-o Èhe

latter group. In adclition its clonal propagation is l.lot effectíve

as in sugar cane or cassava. s and because of self-incompatibilityn it

would be clifficult to fix in the, hotrozygous condition a desj-rable gene

combínation.

From the IÍterature ít would appear that there- are three procecl-

ures in corn breeding which could be applied i:o r¡. Tyegrass, namely

rnass selection, rnass pedigree selection, and synthetic varíety prodrrc-

tion with the constituent genotypes being derived from a mass selecte-d

population.

7 . Micro-envir:onment al variation and the selection of genotypes

Varíous desígns have been suggested to minimlse the effects of

micro-environmental variation on assessme-nts made in mass selection'

These desig¡s are ¡:reant to reduce the masking effect of ttre environment-'

ïhe three most ímportant environmental factors aÏe: soil heterogeneíty,

plant compeËítíon and seasonal variation in the climaLe'

Ideally, an experírnental site should be uniform and clevoi'd of

soil he-terogeneity. Ho\n'ever, in practíce micro-e-nvitonmenüa1 variation

occuïs even in the srnall. areas occupied by selection bl,ocks.

Soí1 heterogeneity causing Don-herÍtable yield differences may

completely mask genetic clifferences. Under such conditioirs Èhe selec-

tÍon of high yielding plants is of no value. Soil heterogeneity is

mosË evident in uniformity trials ,¿here iclentícally Lreatecl plants of

the same genotype do not yíeld equally. Such variation is the cause

of experimental eïror in field exper:iments (Le clerg, 1966). Fí-sher

(f931) has suggestecl that the ef fe-cts of uncontrollable var:iaËion may

t3

be minínrised by replicaEion, ranclomisation and the growing of controls.

To assess micro-ènvíronmental r¡aliation in a sele.ction bloclc, a single

homozygous cultivar, or plauts th4t rqere genetically identícal such

as a clone, ¡eeds to be gro\dl.t as controls throughout the block. In

lü. ryegrass tTris is not possible'

A mettlod which can account for soil heterogeneity in mass selec-

tlon ancl r¿hich may be appropriate Èo t/. ryegrass is selection withín

a subdivided block (Gardner, 1969). Other methods ínclude selection

basecl on a moving meatì and selection based on a response surface

(Hamblín et al., 1977).

I^Iíth regard Èo plant competítion, one approach is to space the

plants at such a distance that competítion does not occur. However,

selection of genotypes groÌ/n under non-competitive situations may have

little value when attempting to improve yield in the competitive

situations of a sward. Studie-s of the yíe1d of genotypes gro!ün under

non-colnpetitive and under svrard conditions showed that the ranking

of the genotypes for yield was ciifferent in the two situations (Knight,

1960). Sel.ection for potential yield per plant based on growing

widely spaced individuals, may not ¡esult in high yielding sv¡ards

(Rogers & Lazenby, L966). With forage yield, highly competitive geno-

types may be desirable, as vigorous genotypes will produce a high

yield of leaves and stems. This sítuatíon may be quite dífferent from

that ín a grain crop. With these, Donald (1968) has suggested selec-

tion should be for genotypes with a low competitive abílity and \tleibe

et al., (1963) consídered that some of the, non-competitive, ge-notypes

are lost in the segregating gene-rations of cereals and that these

genotypes may be high yiel<ling vrtre-n in pure stand.

In Ëhe study of inter-ge-notypic plant corupe-títion in wheat Roy

I{+

(1.g76) state<l that thc most compel-j-tive Êlenotyperì, as shown by survival,

also tended to have a higher ear ntrmber, yield per eaÏ and yield per

plant, showing a continuatiorl of effectil'e co¡tpeti.tj-on into the-

tillering and ear formation stage"

In w. ïyegrass, a continuation of tí11eïing may be regarded as

a desirable character, because it rvill increase the yleld of stems and

leaves. Horvever, in mass selection it is diffi cult to dístinguish

between genotypes rvith a higlr genetíc competitÍve abílity and geno-

types lvhich by chance occupy a favourable rnicro-environnent"

Variation and interactions of genotypes with seasonal climatic

factors such as the flucÈuation of the amourlt and the dís1-::ibution

of raínfall and temperature must be considered as unpredictable en-

vironmental varj-ation. Thís is because v/e cannot predict year-to-year

flucËuatíons and the breedeï cannot deveiop varietíes suiLed to

círcumstances he cannot foresee (Àllard, L964) '

In view of the matters considered in the literature review, it

was decíded:

1. To undertake a study of mass selectíon in w. ryegrass.

2. To grow the plants at a density that I^Iould produce a

sr^¡a::d and ensure comPetitíon.

3. To sub-divide<l the selection bl-ock and thereby selectwithin a small area that hopefully would not show

heterogeitY.

4. To investigate the possibilíty of a breeding procedureinvolving a Progeny fest.

In undertaking this study on mass selection no prejudgement has

been macle on whether a higher yielding, more competítíve IÁI . ryeglass

cultivar is desirable or feasible in Austral.í¿rn agrículÈure. It rvas

undertaken esseutially as a study of the methodolt¡gy of mass selectíon"

15

B}GERII"IIINTAL DETAILS

1. Síte

The experíments \^rere caïriecl out "t th. Irtraite Agr:icultural_t

Research Institute Adelaide, SouÈh Australia (:4o SO South, 13Bo 38t

East, Altitude 120 in). The soíI type vtas å red brown erar:th of the

Urrbrae series (Lítchfield, 1951).

2. Clinate

The region has a mediterranean climate v¡ith cool wet wint.ers and

hot dry summers. The gror.ring season extends through the winter ancl

spring, frorn Apríl/May unËi1 November/December. The average raínfall

for the period Lg25-Ig75 was 629 mm; of which 525 mm (or 83 per cent)

fell duríng the growj.ng season, buÈ the actual amount, in amy one

year can be very variable (Fig. 1). Details of rainfa11, evaporation

and solar radiation are gíven in Table 2.

3 Seed source

The population used in the experiments v¡as developed from a

commercíal supply of seed (ttodgets tr. ryegrass)'

4. Planting Preparation

In 1975 ancl L976,4,350 and 4,537 Jiffy pots respectÍvely were

fí1led with potting soil and used for the gerrnination of the seed.

The seedlings were kept in the glasshouse for thírty days before

transplanting to the field. TransplanÈing was carried out on April 22,

in 1975 and on April 28, ín L976. The procedure of sowing in Jíffy

poÈs and transplanting to the fíe1d was adopted to ensure a complete

stand of plants. Thís \^ras essential for a study of mass selection i.n

a competitíve sítuaÈion.

tI,

F is . I FLUCTUATION IN RAINFALLMEAN MONTHLY RAINFALL IN M]N

130

100

50

19 75

AVERAGES FOR

50 YEARS( 1925 - 1975 )

19?6

tso\

0

J F M A M J J A s o N D

TABLE 2

YearMonth

CLIMATOLOGICAL VAIUES DURTNG THE E)PERIMENT L975-T976

Rainfall in mm and(days with more than I nrrn raín)

7975 r97 6 1925-'75 L975 1976 L925-'75 r975 L976 7925-' 75

Evaporation iu mm

SÈandard A C1ass. Pan.Solar Radiation

ytee I /Yr2 /nay

Jan.Feb.Mar.Aprl.MayJuneJulyAug.Sept.0c È.Nov.Dec.

36. 5))

100.626 .4

110.027 .5

LL9.646 .669 .5

100. 223.26r.0

(s)(1)(1r)(4)(L2)(6)(15)( 13)(r2)(r7)(6)(2)

45.43.62.31.

le)8)14)ls)Is)11)r0)

23.727 .72L.Bs6.881. 573.4Bl .773.56I.L54.r38.430.3

193. 92r4.2L52.2105 .0

67 .244.362. B

59 .492.5

r06.4159.6225.5

2l-3.6206.6161.8109 .8

68.244.058.864.382.4

104. I162.02L2.8

239.200.L7 4.109.63.47.4t.10.96.

r43.r70.2r3.

27.7623.1570"4213 .687.538.317.9L

70.5214.t0l-3.6622.6427.55

27.4623.582r.3213. 039.206.6Õ8.66

70.52l-4.4518. 6925.0027 .65

26 "9224.0219 .3012.93B.t t7 .42-1 1')

10 .3474.6519.6523.7726.04

L7.69.1.

475666

2

6

09

6

2

46

0060

5)7)3)7)e)

((((((((((((

(3.3)(2. e)(3 .3)(7. s)

3

9

6I5

7

I044

7

5

9

(10. s)( ro. 8)(13.6)(12.8)

33.31.

(10.1)(8.e)(s.6)(r1. 7)

!\J

Year 668.4 (104) s1s.1 (123) 629.0 (94.0) 1403.0 1495.8 1s80.4 16.33 17.18 L6.76

1B

5. PLant s crn

The plant spacing was 12.5 c.m x 12.5 cm to give a plant clensiÈy

or. 64 plants/rn2.

6 Fertilizing

The fertiLizers used vlere:

a. Urea

b. Superphosphate

c. Muriate Potash

tl.6 gram/rn2 (462 N)

26.0 gratt/n2 (8.6"Á P)

6.5 gram/mz (50% K)

Applícations \^/ere made at three tímes; after transplanting and

after the fírst and second cutting. The above rate I^tas applied aÈ

each application.

7. Harvest

PlanÈs were harvested three times by cutting Èo a uniform height

of 4 crn above ground level. The first harvest hTas taken about thirty

days after transplanting to the fÍeld. subseque-nÈ1y the intervals

between cuts was four to fíve weeks (fatte 3).

TABLE 3 THE DATE OF TRANSPLANTING AND HARVESTING

Year Transplanting 1st Harvest 2nd Harvest 3rd Harvest

t975r976

22 - 23lhpriI28 - 29/ÃpriL

27 - 29h[ay2 - 4/June

25 / J:u¡:ne

30/June30 - 3l/July28 - 29/Jv,Iy

24-29-

B. Yield

At harvest the forage from each plant was placed in a numbered

paper bag. The number indicated the position of the plant in the

fíeld layout. The material was placecl in a dryiug oven at 70oC, for

al

19

30 hours. In the results, yÍ.eld is e>'pressed as dIy TnatteÏ per

plant-. The detaj-led yields aïe presented in appendices. Ttre

yields are presented in the. same order as Èhe plants occuTred in the

field platrting and are the total.s of the three cuts for Ëhe consecutive

trials.

9. Field layout

The field layours for 1975 a:rd 1976 al.e gi'ven in Fig. 2 and

3. Two rows around the plots or blocks rn'ere used as borders. In

management, they were Èreated identically as the tesÈ plants (e.g.

fertilizer and harvestÍng) buË the results are rlot considered'

10. The selectign sequence

In 1975, Ëhe maín plots (numbered 1-16) did not represent dif-

ferent treatments. They are. referred to as plots because in the

second year rnain plots of the same size were used. In 1975 sixteen

plots of 196 plant per p1ot, formj-ng a population of 3,136 plants,

were gro\^/n. Each pl.ot was divided into four sub-plots, providing

49 plants in each sub-plot. The three highest yíeldíng plants,

based on the total yield of the three harvests v¡ere selected from

each sub-p1ot, resulting in L92 selected plants. These selected

plants kTere transplante<1 after the third harvest to plastic pots to

allow them to interpollinate withín a glasshouse. Out of 192 selected

plants 188 survived the tra-nsplanting while the other four died before

setting seed. In order to produce seJ-f-pollinated seed, three to

fíve tíllers \,rere bagged on each p1ant. Cross-pollinated seed was

obtained from tíllers that \./e-re not bagged. fn December, 1975, Lhe

cross-pollinated seed and self-pollinate<l bags we,re harvested separately

from each p1ant.

2-O

d,tiI

LAY GUT ffiF T þI E ËX PER Iru¡Ë hITI N 197 5

3 PLANTS SELECTED FROM EACH SUB PLOT OF 19 PLANTS ON

A BASIS OF TOTAL YIELD.

Íis-2

I

t

1 2

5 6

10I

14f3

1211

1615

43

B7

oeo@c e)o co oc 'o e o o

C oC O

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o

e

6

e

eo

ø

C

o

e

oo

oo

o

e

c

OaOOC¡lÐOOOoeCe

@t@". cO.Ð.ocoêoaGooo6C CO O O O O

aoocca9

Ê

o

co

eo

o

oocaCOOCOO9Ooo

ooGE

@o99oCGOeo

OOoooO,@OOaa

@o

o|D6('

¡O

aOOOOOOOCègO oc o

c@ " c e o o

eoooêoc@o . ' o o o

o o . o 6 e@

2-1.

rl

L

I

(

tt

I

t,tr

l

lrII

LAY OUT OF THE EXPE$ÌIMENTlN 197 6

aloao@aoao

aco@oaoaaooaao

@..aaaooa.oooo o

oooooa¡oao

ooocoocoo@.'ooa

oaGCaooooo

aooaOOOao¡r

ooc n@@"ôcaoceo

oeooaaoo

aooooooaoot

o oa

aCooaO¡Ocaoocc

ao

@@a

oaaooooc

ooaoaoooac@o o o@.Oo o.OCOaOoOOoa

caoooooaaaaaoooaooa

3 PLANTS SELECT E D FROM EACH SUBPLOT OF 19 PLANTS ON

A BASIS OF TOTAL YIELD.

r

12

5 6

I 10

13 14

17 18

4J

I7

1211

1615

2019

F is.3

22

Tn I976, the sowíng and transpl-anting proccdure was repeated

but the number of plots was incr:eased by four. Twenty plants origína-

tíng from the seed of each selected plant \^7ere gror¡rrl' 'rhe population

in thís second cycle consisted. of 3,92.0 plants (1BB x 20 plants and

160 plants from seed of Ëhe original or unselected population) ' In

each plot Èhere were 1BB and B plants' gro\^7n at::andom' Thís procedure

is similar to eaï to ro\,ù sel.ection in corn, with the difference Èhat

here each plant was planÈed at randotn as an individual in a plot, not'

in a row system.

A summary of procedures is given below:

Tn the field In the qlasshouse

Initíal populaÈion3,136 plants

r975

192 plants ü/ereselected

3 to 5 tillers on eachplant were bagged andthe resÈ left oPen tofacilitate intercrossing

J20 plants from oPenpollinated seeds of eachof 1BB selected plants,and 160 plants from seedof original poPulaÈion

3 to 5 tillers from eachplant were bagged and therest left open to facili-tate intercrossing.2 to 3 tillers fromdifferent plants werebagged to check seed setunder bag.

__>

Second cycle3,920 planÈs

r976

240 plants vlereselected

__Þ

i

23

I

RESULTS

The objecÈíve of this experime-nt on w. ryegrass vias to study the

effect of mass selectíon for yie.ld. Addltional objectj.ves \^lere Èo

investi.gate the possíbility of producing se1.f-pollinated seed and to

select individual plants rvhích coul.d be used as a constituent in a

synthetic variety.

1. Yield results

The yiel-d \^,as measured as total dry mat.Èer obtained from three

cuttings made at inÈervals of approxi.mately 30 days. An overall

impression of the resulËs for the tr^7o years may be- obÈaíned from the

hístograms and statistics in tr'ig. 4,5, 6 and 7. These hístograms

of yield per plant at each cut and the total of the three cuts are

gíven for the consecutíve years. The histograms l^lere obtaíned by

dividing the range of the yield into fifteen classes and calculating

the pergentage in each class. The statisEical values show:

a. Statistics relevant to'the fi::st year

i A wide range in yields l^7as present one month aftertransplanting. This range increased with successíveharvests.

1i The large plants at the first harvest continued to belarge. This was evident for many plants but it can beseen for the plant with the bighest total- yield forthe year. It had individual cut yields that \'{ere nearthe híghest, for each cut (total = 22.7; individualyields in each cut 5.6, 4.8 and I2.3).

iíi. Skewed distributions were obtained. The skew íncreasedprogressively wíth each cut.

At the end of the first year some plants died in the field after the

thlrd harvest. However, it díd not affect the sele-ction because most

of the deaths occurred among the weaker plants and only a fer¿ of the-

highest yieldíng plants were affected.

i

24

No. of Plants 3136Min. .10Max. 5.60Mean 2.127SE (Mean) .0q131Variance .5373c.v. .3446Skewness .5844SE (Skew) .0437Kurtosis .7399SE (t<urt) .087 4

Fígure 4alsÈ Harvest 27-28 /5/L97s

No. of Plants 3136l"lín. . 10Max. 5.30Mean 2.048SE (Mean) .OL32Variance .5427c.v. .3597Skev¡ness .6118SE (Skew) .0437KurÈosís .6420SE (Kurt) .0874

Fleure 4b2nd Harvest 24-25 /6/L975

'No. of Plants 3136Min. .10Max. 13.60lulean 3.931SE (Mean) .0383Variance 4.59L5c.v. .5451-Skewness .8222SE (Skew) ,0437KurËosis .7573SE (Kurt) .0874

Fígure 4crd HarvesÈ 30-31 7 /Le1s

21

AF

R

E

aU

E

N

c

l0

t8

t2

o

t8

l2

l8

tz

N

Io

2{

o I z 3 ¡t 5 5 ? I 9 lo ll 12 13 Y¡eld lt'

O I ¿ 3 I 5 6 7 I s lO ll 12 13 Y¡eld s

F

R

E

aU

E

N

cY

I

N

olo

6

o

z1

F

R

E

aU

E

N

cY

I

N

"1"

9123{ 5 o 7 E M ll t2 13 Yields

B

c

o

25

No. of Plants 3920Min. .10Max. 4.00Mean 1.820SE (Mean) .0092Varíance .3328Skewness .3LI2SE (Skew) .0391Kurtosis .3028SE (Kurt) "0782

Fígure 5alst llarve sx 2-4/ 6 / L976

No. of Plants 3920Min. .10Max. 6.20Mean 2.247SE (Mean) .II22Variance .4937c.v. . 3128Skewness .4555SE (Skew) .0391KurÈosis .5903SE (Kurt) .0782

TÍgure 5b2nd Harvest 29-30 / 6 / 7976

No. of Plants 3920Min. .10Max. 6.30Mean I.929SE (t"tean) .IL22Varlance .7937c.v. .3128Skewness .6474SE (skew)Kurtosis .8163SE (Kurt) .0782

Fígure 5c3rd Harves:u 28-29 / 7 /L97 6

21

F

R

E

oU

É.

N

cY

t8

t2

I

N

7"

0

2l

0 | 2 3 4 5 6 7 8 I Y¡eld s

0 | 2 3 1 5 6 7 I 9 Yields

0 I 2 g 1 5 6 7 I 9 Yieldg

F

R

E

aU

E

N

cY

I

N

%

I

N

t8

t2

0

l8

l2

24

F

R

E

oU

E

N

cY

6

A

B

c

7"

0

26

Nr:. of PlanÈs 3136Ivlin. . 1.0

Max. 22.70Mean 8.106SE (Mean) .0573Varíance L0.2970c. v. .3959Skewness ,6816SE (Skew) .0437Kurtosis .5946SE (Kurt) .OB74

Tigure 6aTotal Harvest 1975

No. of Plants 3920Mín. .10Max. 15.60Mean 5.996SE (Mean) .0284Varíance 3.772c.v. .2970Skewne.ss .3195SE (Skeq¡) .0391Kurtosi s .6794SE (Kurt) .0782

Fígure 6bTotal Harvest 1976

6

îR

E

aU

E

N

cY

I

N

T"

F

R

E

ou

E

N

cY

I

N

%

24

18

',2

24

18

12

6

0

I 975

0 6 E t0 12 14 f6 lB Z0 22 yrELD s24

0

20 4 6 I l0 12 14 16 18 20 22

1

27

.Na. of PlanÈs 160Mln. 1.90Max. 13.60Mean 5.303SE (Mean) .1380Variance 3.0454c.v. 3.29rSker^¡ness L.292OSE (skew) .1918KurÈosis 3.0197SE (Kurt) .3814

Iieure 7aOrígínal Population 1976

No. of Plants 3760Min. .10Max. 15.60Mean 6.026SE (Mean) .0289Varíance 3.1568c. v. .2949Skewness .3474SE (Skew) .4806Kurtosis .4806SE (Kurt) .0784

Figure 7blst Mass Selected Population 1976

A

0

B

27

I

21

r5

2l

f5

tR

E

ou

E

N

cY

I

N

%3

l2 11 l6 Yield g.

te t1 l6 Y¡eld s.

I l0

8

2 10

27

F

R

E

oU

E

N

cY

t0N

Io3

0

0I20

¿()

b. Statístics relevant to the second year (1976)

The points made in i-iíi above were also relevant to the second

year. Adclitional asPects are:

iv Yielcls Í-n general were l-ower than in the firstyear (c.f . Fig. 5 with FÍ-g" 4 or see Fig' 6) '

v The yields of the selected population vr'as greaterthan the urrselecÈed cont::ol population (Fig' 7) '

vi The slcew of ttre seiected popul-ation was less thanthe slcew of the unselectecl population (statisticspresented with Fig. 7).

Reference will be made to some other aspects of these results. I¡trhen

the 1976 trial is considerecl as one population the results show a

similar trend to the first year wíth the skewness of the distrí.butions

tenclíng to increase with each cut. Total yíelds were much lower in

1976 than ín L975 (¡'g.. 6a and b); a díffereflce that is probably

aËtributable to rainfa1l" Irrom the clinatoloqical values presented in

Fíg. 1, it. is evident that the rainfall distributÍon in the two con-

secutíve years de-viatecl sigrrificantly from the average for fifty

years; in Lg75 it was above average and in 1976 below the- avelage'

As 1976 \^/as so dry it is understandable that the yíelds of both the

selected and unselecÈed populations \dere less'

Because the environments in the consecutive years \^7ere so diffe-r-

ent the best measure of the effect of selection can be obtaíned from

Lhe comparison between the selectecl and unselected (control) population

grov/n in the same yeaï (L976). The statistical values (Fig. 7) show

that the mean yield of the selected populati-on in 1976 was 13'B per

cent greater than the mean yield of the contr:ol. Total yíe1ds are

gÍ.ven as the value for e.ach popuJ-atíon on a pe.r plant and per hectare

basis (tab1e 4E and b).

29

TAtsLE 4A THE MEAN OF TTIB TOTAI, YIELD FOR TIIREE CUTTINGS

G/PLANT DRY }ÍATTER

Year OriginalPopulation

1st cycle of masssele-cted poptrlation

È

t97s

r976

8.10 r .06

5.30 1 ,74 6.03 t .03 5.046^')kr(

?k?kt< highly signfficant at 0.1% 1evel (tt"b 3.29r)

TABLE 4b TOTAL YIELD FROM THREE CUTTINGS IN KG/HA DRY MATTER

PLANT DENSITY 640 000

Year OriginalPopulatíon

lst cycle of massselecËed population

L975

r976

52L6

3392 3840

These total yields should noÈ be compared directly with published

annual yíelds of w. ryegrass, as the forage produced after the third

harvest ín the present trials ís not included in the yielcl calculation.

Table 4 represents yields up to the begínning of August when- the

selected plants r{ere moved to the glasshouse.

Because of the skewed nature of the popul-ations it was decided

to make a test, addÍtíona1 to the one in Table 4a, for the sígnifÍcance

of difference beËween selected and unselected populatj-ons.

Calculation of Èhe significance based on a logariihruic arrd square

root transformation of the data did not alLer the conclusion that

Èhere \ras a very highly signÍ ficanE dif ference betr¿ee¡r the- selected

and unselected populatíon in 1976.

(t. = 4.76I and t = 5.117)' Iog sqrf.

30

c. Herital-:ílity

An esËimate of the heritabili.ty of yield was obtained for this)

population of w. ryegrass. Heritabi.lity (h-) is normally calculated

AS

where,

R (response) is Èhe difference between the means of the selected and

unselectecl population, and would be, in this study the dífferenc'e

between the mean of selected population in 1976 and Èhe original

populaÈion in 1975.

S (selection differential) is the dífference between the mean of the

selecËed oï truncated planÈs ín Èhe original population and the me'an

of that oríginal populatlon. With Èhe normal method of calculation

the S value ín this study would be derived from 1975 data.

Because of Èhe large environmental dífference betwee-n years, and

the resultant lower yields in the second year Èhis normal method of

calculating heritabilÍty is me.aningless. For instance, R ís negative

(see Table 5). However, a heritability calculation vras made usí1g

an R derived from the difference between the mean of the selected popu-

lation when grown ín 1976 and the original population grown in 1976

(control) .

Two S val-ues were derived. one was that normally used and

described for S above. The second S value \'¡as calculated by obtaining

the díffeïence between the mean of the highest yielding 6 per cent of

the conËrol planËs in 1976 and the mean of the conÈrol popul-atíon in

Lg76. The results of these calculaLíons aÏe given in Table 5.

:,zRS

31

TABLE 5 THE HERTTABTT,TTY OI TlrE YTELD 0F I^1. RYEGRASS

IN THE lST I"ÍASS SELECT]OT{

Population Mean ofPopulation

Mean of 6%

TruncationSelecË1onDífferential

(s)

Response(R)

Original 1975

Original 1976

Selected 1976

B. 10

5 .30

6.03

14. 38

B. B8

6.28

3.58

6"03-8.10=-1.076.03-5.30=0.73

trlhen S of 1975 ís used:

Heritabilíty = 0-:_73

6.28.1r.6

or

S of 1976 is used:

Heritabílíty = 0.73 = .2O43.58

2. Affect of select ion on t a ç 1¿ er^r o f the nonrrlation

trlhen a comparison is made of the total yíeld distributÍon of the

mass selected population ín L976, it is evident that íts skewness i.s

less than both the unselected populatíons in 1975 and 1976 respectj,vely

(Fig. 7b, 7a and 6a). Furthermo::e the sker¿ fqr the unselected popula-

Èion ín 1975 is less than the skew for the unselected population in

r976.

A rneasure of the affect of selection may be obËained by calcul-a-

Èing the percentage of plant.s above or below the mean yield of the

population. Thís calculatíon is based on the assumptfon that if the

reactíon of every genotype ín the populatíorr to the different environ-

ments is linear or pïoportionate, the percenËage of the genotypes below

and above the mean of the population should be unchanged. Àn alterna-

tive cal.culatlon was based on pelcentages below and above the mode.

The purpose of these calculations \^ras to take aceount of the skew and

32

see whether aS a tesult of lnåcs seleulíon thele rl¡as êny changel in

these percentages. The results are. given in Table 6a and b.

TABLE 6A TFIE PERCBNTAGE OF ]]HE PLANTS RELOW AND ASOVE

THE I"ßAN OF THB POPULATION

Year Population PopulatíonMean Yield

Plant Percentagebelow rnean above mean

r975

L97 6

L97 6

UnselecÈed

Unselected

Selectecl

8.10 t 0.06

5.30 I 0.03

6.O2 ! 0.L4

55.20

58. 75

53.09

44. B0

4L.25

46.9r

TABLE 6b TIIE PERCENTAGE O}- THE PLANTS BELOI.I ABOVE AND

AT THE MODE OF THE POPULATION

Year Population PopulationMode Yield

Plant Percentaqebelow mode aÉ mode above mode

L975

r976

r976

Unselected

Unselected

Selected

6.13

4.24

5.27

29.76

28 .38

34.58

L9.29

24.38

22.47

50.95

47.49

42.95

TABLE 7 THE PERCENTAGE OF TIIE PLANTS BELOW

THE I'IODE AND TFIE MEAN AND T'HE

SI(E.WNESS OF THE POPULATION

Year Populatíon P1.ant 7"

Below theMode

PLanE 7"

beloi^r ÈheMean

Skewness

r975

r976

r976

Unselected

Unselected

Selected

29.76

28. 38

34.58

55.20

58.75

53. 09

.6818

L.2920

.347 4

33

TABI,E B THE VALUES O}' MODE l"tlIAN ANn SKEhTNESS

OF Tl{E POPUI,ATION

Year Populatíon Mode Mean Slcewness

L975

L976

r976

Unselected

Unselected

Se,lected

6 .13

4.24

5.27

B. 106

5.303

6.026

.6818

r.2920.347 4

From Table 6, iÈ can be seen that both the environme'nt and mass

selectíon caused a change in the- percentage of plants below and above

the mean and mode of the population. The change in percenËages of the

plants belor¿ Ëhe mean and mode ís consistent with the values of the

ske\nrness of the populaÈion (Table 7).

3. Environmental hetero seneity with in the experimental areas

The size of the experimental areas for the trials was 8.75 rn x

8.75 rn in 1975 and 8"75 rn x 9.50 m ín 1976. By plotting the mean

yield of each sub-plot, it can be seen that even such smal1 areas show

variation in the environment (Iig. 8). Tn 1975 yield Per square meter

varied from 6.91 to 8.87 ancl in 1976 from 5.00 to 6.75 g. The grouping

of high yíelding and lov¡ yielding plots suggested that the micro-

environmental varlation was not random. The.se tr^/o matters' of large

micro-environment-al varÍatíon and íts non-randonmess indicate the

need for undertaking mass selection on a subdivided plot basis '

A comparíson of the plants thât v¡ould have been selected if tnere

had been no subdivísion of the total block, and the plants tÏraÈ

actually were selected is presented in the next sectj-on.

3t:

MI CROENVIRONM TNTAL VAR IAT ION.TH E MEAN Y ¡ELDS OF TI.{E 49 PLANTS IN EACH SUBPLOT.

POPULATTON MEAN YTELD I.l01975 T'R E AL

LESS THAId 8.OO

8.00 - B .50 F¡s. 8 ar-ll-Jl-J

ffi

9.13

- E .3',1 7.91 .07

7.89 11 7.87 I 7.5805

7.97 7.s2 $ 7.73 7.Tg

7.89 8.1 7 .7s 7. s I 7.58 7"50 l'At

5 7.69 6.9f 9.24 -:8"2 7.84 7"867.77

:8.13=

8.14

F-s.sz#

{-H+t+

tì7

8.68 (}t! 5 8.4î: 8.ü3

= 8.16-

6 ?.8{} 7.8S

7.97 7.78 8.7?

_Q ã Iîæ(}i-P.crd (l ûr¡

"¡

EarJ 7.9S

&.4

I 5 u ff I A(ò =së=Ëiüs.74cl ç?

å,, . t.tt

ABOV E 8.5 0

35

POPULATION MEAN YIELD

M ICRO LNVIRONMENl'AL VA l{l Al lO¡,¡.

THE MËAN YIË.LDS OF TI-IE 49 PLANTS IN IACH SUBPLOT.

5:9 IX976 T'REAL

LËSS THAN 5.69

trffi

5.70 - 6.39 Fis.8 B

5.6 35.5 0 1* 0.7 4 6.6 S

6.tS-- 6. I 6.585.90-__- 5 ì.] "gJ ; 6.t6-++6.4t1+-I

ffi

5.6 0 s_ 5 o(, 0 þ tÐ 7 I s I I 5ôt) ?tt t 0 5

5.33 5.60 6.28:6.02== 6.13 --6

11 .13 5.63

5.11 0J:, î 5 6 n1 h z 't:5'S 5.61

6.11 {i,ü2 6.38-:S.0? 6.6 5.7 5.5 ?

5-8 3=- 6.27 =::5.6a: 5.99-- 5 .98 7$ì- 5.67

5,9 3--_ 6.0 5.63 6.20 - 6.X1: 5"?3= 5.91'=-5"$2

5.3 3 6.16 -

{j s5" 8,Ê---* ri. --- 6.2

5"4õ

5.65

6.68 5.t 05.{}2.--_ 6.39.-- 6.32 _:6 '86 -- 6. 10

ABOVE 6 .39

36

4. The d istribuLiurrs of vield in popr-rlat ions of massl selected T¡lanto

Both clisfributions are basefl on the supposition that 6 per c.ent

of the population was ::etained (Fíg. 9 and 10). Understandably the

range of Èhe selecte<l plants from Lhe subdivlded plots lvas much greater

than if simple truncation of the whole population had been used. Some

of the plants selected on a basis of the subdivided ploÈ were from

loW yielcling areas and were only L2.5 per cent above the mean of the

population compared with 62.5 per cent above the mean for: the lowest

yíeldíng planË selected under fu1l truncation.

5. Proge-ny test in r,¡. ryegrass

The prqgeny test to be presented used as arr evaluation, the

frequency with which the offspring occurred among the highest yie-1díng

plants in their generation. Each of thê lBB selected plants in 1975

\¡ras represented by twenty offspring in 1976' randomized over twenty

plots. Thus each plot contalned 188 different offspring. In addi.tion'

there wele eight unselected plants (original population) as controls.

In total, each plot contained 196 plants. To be comparable to the 1975

trial the 196 plant plots were divided into four sub-plots.

An evaluation was carried out based on the best three plants in

each sub-plot. If the t\,renty offspring of a selection \^/ere among the

best ín each p1-ot the selectÍon v¡ou1d have a score of twenty and the

selection could be regarded as having a high combíning ability. On

the other hand íf the selected planÈts offspríng occurred infrequently

the selecte-d plants may have had a::elative high yield in I975, the

year of selectiorì, as a result of specific combining abilíty or because

it was gro\^rn in a favourable positior-r (i.u. an envitc¡nmental effect).

Using the subdivided plot procedr¡r:e reduces the likelíhood that the

iÈ.T

I

37

DistïibuÈíon of selecÈed plants lfselection had been based on tTuca-tion of the whole PoPulation and no

account taken of subPlots (67" of

fopulatíon selected, 192 Plants) 'ir"rrg. L3-22 gram. The dotted Partof ttre histogram above the classlnterval centred on l3g indicatesthat only a proportion of the plantsin this ctrass were needed to make uP

6iÁ of the PoPulaËíon

Figure 9ar975

Distribution of selected PlanÈsbased on subclivided plot truncation$% of. poPulation selected, I92plants). Range 9-22 gtams'

tr'ísure 9b

T

rl

L975

100

60

80 AN

o

oIr

P

L.4.

ÌiT

0

60

40

20

40

20

10 13 16 19 22

YI.ELD IN GP"A}.IS

10 13 16 19 22

BN

o

oF

P

I,A

ryT

0

YIlji,D n{ (]lì¡ir.s

I

i

l

I

I

I

3B

Distributíon of selecÈed Plants ifselection had been based on trunca-tion of Èhe whole PoPulation and no

account taken of subPlots (6% ofpopulatíon selected, 240 Plants) 'Range 9-15 gram. The dotEed Partof the histogram above the classlnterval centred on 9g. indicatesÈhaË only a proportion of the plantsín thís class were needed to make uP

6% of the PoPulation.

Fieure 10aL976

DlsÈríbution of selected plants basedon subdívided plot truncatíon. (67"

of population selected, 240 plants) '

Aa

Ia

I

Ë--

150

130

80

60

40

20

N

o

oF

P

L¿I

I'l

ll

8 f0 12\4 16

YIE],D I}i GRAì.IS

8 10 12 ttYII:LD Iii CI,^ì S

6

B80

0

100

60

40

20

I\¡

o

oF

P

LAN

T

06

I

l

l

I

I

Ii

I

16

39

selection was high yieldi.ng because of a favorrrable environment' The

daËa are presented in Table 9. The best selec.Lion (No. 11) had 10

out of its 20 offspring anìong the highest yieldíng plants in each ploÈ'

Only three others had a score of 6 out of 2O '

T'he parents and offspring could be ranked in seve-ral rvays to

illustrate the relation between them. They could be. ranked:

a. On a basis of the yield of the parenËs'

b. On a basis of the frequency of reselection of theoffspring.

c.onabasisoftheir:offspríngmeanyield(Tab1e10c).

Tables lOa and b contain the fírst of these two rankings. The third

is not pres.ot.d as it can be readíly derived from Table 10b ' By

using these types of ranking, there are three possibílities for

selectÍng plants for compounding Èo form a synthetic variety' However,

which one r¿ould give the highest yield has not been tested. It has been

suggested that the optimum number of constituerìts for diploids lies

between four ancl ten (Gallais eË aI., 1970) '

some point relating to these proceduÏes aÏe referred to in the

dis cussíon.

6 Self-pollinated seed

To determine if ít would be possible to produce self=pollinated

seed, three to five tillers of Èhe lBB plants in 1975 and 24O plants

in 1976, were self-po11ínated by bagging with glassine paper bags in

the glasshouse. Not a single plant p:oduced any seed. Thís l-ack of

seed could have been due to self-sterí1ity or to conditions wíthi-n the

bags that \^rere unsuitable for seecl set. The g1.ass:Lne bags were those

used customarily for |solation in the pollínation of rtheat, barley and

other grasses in the glasshouse and there \'Ias no ínítial reason to

40

TABLE 9 THE FREQTIENCY I^IT TH I,/}TI.CH À SEI,ECTED II,ANT'S OFFSPRING

OCCURS A¡4ONG TIi Ïi TlIGHEST YIELDINGP'IÁNTS IN TI]E N]]XT G]]NERÀiT].ON

I'OSSTBLE S 20

or.rL. F.R. s.pL. F.R. s.PL. F.R. s.PL. F.R" s.PL' F'R' s'PL' F'R'No. No. No. No. No. No.

1120

154].62101115r49lB3

350

L43159170

4L21323243645596669739799

100106113TL4148168

155L63L72LB2

2

9

15t7222529323B40424951.

5860677779B1B5

B6

L44l'5715B160161r66L69172r73L74175t7BL79185lBB

56

I

L46r52r 53155r56]-57I7IL 74775r16178180184r87188091o920960 980 1030 1.04

0 1080 ll10 lLz0 1170 1180 1190 1200 121o L22o r23o r2.4

L25l-27r28'r29

131133135L36r3B139L42L47150151153L56165L67L7L]-76180IB4186TB7189190191L92

000000000000000000000000000

:

93

222

222

2222222222222222

22222

2

22

22

I7

10

106

6

6

55

55444443333333333333333333

1B2627354748546I636465687072B3

899095

L02107110116126130L32140L45L46l-52

2

222111tI1111II1I11I111I111111II

00000000000000000000000000000000

L4

2B

16192L

94105109134L37L47

JU31333437394L4344

* S.Pl. No.

F.R

A selected plant íderrÈífication number

Frequency of ::eselection i.e. selectecl again

4t

TABLE 10a A RANKTNG OT SE LECTED PI.ANTS ON A BÀSIS OI¡ .'ry!.18"YIELD TOGE]]HER I\ÌlTll TIJE FBEQ UENCY OIT RESELEC]'i_a{

OF OFFSPRINT.} AND TI{E 20 OFITSPRING }181N YIEI,D

Selected YieldPlantNo.

F. R. Of f sprÍ.ngMeanYield

Sele-c.Èed Yie-ldPlanÈNo. .

F.R. OffspringMeanYi eld

t6490B4BB

894s

116856775

115L75

385113

15053

LB25B30373210

L43168

932B

100140159t67

703

98L76

1439

12LL73183172

4920B]42

LL4

6.425.124.796.087,L26.2L6.s16.285 .827.385.156 .356 .466.825.625.286.9L5 .836.7L5.746.Bs6.006.996.535.7s6.s37.LB6 .816.995 .896,r26.526.L25 .6s5.274.825 .486 .316.696.306 .557.3r4.956.386.52

50104186lBB

46r01134]-62l.1B166

4LLs7

77TI2

2572

L54B691

L02141

7999

111L24127L49L79

4757

r2026

135156L72

2735

rt7r30151170153l-74

BO

r04155

6.425 .11lr.9B6.345 .586 .604.977.L35.7?-5.825.626.576.L76.486.10s.677 .805 .455.9s6.1r5,326.606 .995.675.335.827 .495.575.975 .34s .986.BB6.r75 .077 .066.235 .406 "797 .O25.627 .66Ê 1a)"lL

6.?-55. B0

6.2.86 .50

2

002

32

31051II3002

I0012

43I032

402

40000015II60I3

22.720.7t9.919 ,619 .5L9.219 .018. B

18.61B .518 .518 .5t8 .418.418.318.318 .118 .117 .7l-7.5L7 .517.117 .0]-6.9L6.916 .8L6.716.7L6.7L6.7L6.7L6.616 .516 .516 .516 .316 .316 .316 .3L6.216 .116 .015 .915 .915"8ls. B

L5.7 4

r5.7 0r5.7 0L5.7 1

15.6 015.6 5

15.6 1

15.6 6

15.5 015.5 1

15.3 015.3 I15.1 115.1 015.0 115.0 2

15.0 6

14.9 I14.9 01.4.9 2

L4.9 I14.8 1

14.8 3

14.8 014.8 014.8 014.8 514.8 Ir4.7 2

L4.7 0L4.7 014.6 2

14.6 0L4.6 0].4.6 2

14.5 214.5 2

L4.5 0L4.5 2

L4"5 014.5 4r4.4 0L4.4 I1.4.3 014.3 0L4.3 2

42

TAIILE 10a (Continued)

Se-lected YieldPlantNo.

F.R. Offsirringl'{eanYiel-d

Sele-cted YieldPlantNo.

F.R. OffspríngMeanYíe1d

5.926.975 .68

6.2L5 .815 .985.456.745.776.374.695 "77s .386.2L6.046.806.065 .586.28

13.0 013.0 2

l-3.0 013.072.9 1r2.9 1L2.9 0r2.9 1L2.9 3

Lz.B O

12.8 2

I2.B O

r2.8 0

7B

L22r26133

,(1812260

4.926.736.395 .835 .006.595 .046.696 .586.275.525.826 .86s .606 .405 .355.476.345.515.665.525.286 .086 .837 .995 .185. 135.235 .83s.887 .035.525.726.L44.875 .546.O25. 706 .604.936 "004.Bs6.145.794.825.625 .435 .806 "046.93

14.3 0Lt+.z 3r4.2 2

r4.2 1

14.r 0

14.r 2

r4.0 014.0 3

14.0 2

14.0 314.0 113.9 113.9 3

r3.9 I13.8 1

13.8 0l3.B 0L3.7 2

L3.6 0

13.6 1

13.6 013.6 013.6 2

13.6 2

13.5 1013.s 013.5 013.5 013.5 I13.5 113.4 3

r3.4 0

l-3.4 Ir3.3 2

r3.3 013.3 013.3 013.3 113.3 2

r3.2 0r3.2 IL3.2 013.1 0r3"1 0r3.l 013.1 013.0 013.0 013.0 013.0 3

189148L52T7L

3395525965

106r37

9497

161160180187

5431404476

107L32

117L7492

158L69

24L42185

183456

103r091r0

96144t92'19108L2B131

162L4369

BI113

683

l-25136L7Bl5

tL729

7

5561

*62*82105

t64

]-29L4s

9

366368

139119138163

5

48T266

1912

234

L47B

L23165146

73190

5.946.426 .9ss.006.494.926.565 .415 .575 .615.254.976.L25.165.586.506 .805 .835 .906.735. 895.785.205.sB6.r46.266.4L6 .47

L2.B 1L2.7 IL2.7 Ir2.7 112.6 2

L2.6 0L2.6 2

12.6

L2.4 2L2.4 0L2.4 2

r2.3 Ir2.3 3

L2.3 2

1.2.3 2

Lz.T O

I2.L 2

12.0 0r2.o 2

11.9 311.9 311.9 011.7 1r1 .7 3

1r.6 311.5 011.3 011 .3 011.3 0

12.6I2.5 1

r2.4 2

]-2.2 0l-z.L O

11.0 2

i0.9 39.9 0

/+3

TABLE 1Ob A RANK.I NG OF TI1]Ì SIlI,IÌC TED PLANTS tsASE]] ON }'REQUENCY

ol' Rlrls!il,liU'lrI0N uF UFFSPR].¡IG TOGET}IER I.JI'I'iI

THIIR YIiiLD Al'lD 20 OIIFSPRING I"J.EAN YIEIJ)

Selected YieldPlantNo.

F. R. Of f spr:ingMeanYield.

Sel.e-cted YieldPlantNo.

F .l{. 0f f springIleanYield

l120

154L6210111.5r49183

350

L43159170

4T2132324364s596669739799

10010611311414816B

17

1018262735474B5461636l+

65

7 .997.3r7 .807.r36. 607 "387 .496.696.526 .426.996.997 .66s.B95.406.826.737 .036.56t.r26.696 .806.936.4r6 .686.997 .186.276.746.526.736..536.476 .066.006.L46.BB6.23s. 405.975.586.346.285.416.956 .58

6B7072B3899095

r02107110116L26130l-32140r45L46L52155163L77782

2

9151722252910JL

38404249515B606l7779B1gs

B69394

r05

5.s76.L25.676.376 .086.476.596 .116 .086 .606,216.917.026 .836.Br6.496 "266.396 .506.r27 .066 .91s .904.926.2L6.O46.2L6.106 .806. B56.355.666.386 .556 .465. B35 .816.286.L76 .605 .456.515.455.75_5.825.9/1

t2.3 2

16.6 2

15.0 2

T2.B 2

19.5 2

20.7 2

14.1 2

L4.9 2-

13.6 '2

13.3 2

19.0 2

r3.0 2

L4.5 2

13.6 2

L6.7 2

r2.4 2

11.0 2

r4.2 2

1.4.3 2

I2.I 2

l.4.6 2

18.1 2

r1_.7 l.L2.3 1

L2.7 1

r2.7 1L2.9 1

15.0 1l_2.7 I17.1 118.4 1

13.6 I15.8 I16.0 1IB.4 117.7 IL2.9 I18.6 I15.1 114.8 1

L2.9 I10.8 1

L4.9 116 .8 1.

13.9 112.5 I

13.5 l015.9 6

15.0 6

15.6 6

15.6 5

18.5 5

14.8 5

16.2 5

16.5 4

r5,7 416.9 4

!6.7 4

14.5 411.6 3

1r.9 318.3 3

TL.7 3

r3.4 3

L2.3 3

L9.2 314.0 3

11.9 313.0 3

10.9 3

13.9 3

14.8 3

16.7 3

r4.0 3

L2,9 315.9 3

L4.2 3

L6.9 3

L2.4 2

L2"6 2

17.0 2

13.3 2

L4.6 2

L4.5 2

14.5 2

r4.7 2

Lz.O 2

L3.2 2

L2.6 2

L2.3 2

L2.4 2

14.0 2

44

TA3I.E 1Ob Continued

Selected YieldPlanÈNo.

F,R. 0ffspringluleanYie-ld

Selected YieldPlant I'

No.

F.R. 0f f s;pringMeanYíelcl

109134r3714rL44L57158160161l66169t72L73t74n5178t79185lBB

56B

L416192r'283031333437394L43444652535556577L7475767BBO

5.704.975.525.326 .006.57s.B36 .405 .605.825 .886.306 .316.255.155.385.575.746.345 .16s.775.205,27s .436.745 .806.046.7L5.575 .004.875.744.825.646 .045.525 .585 .045.285 .585 ,545 .345 .185"135 .82s.2B5.985 .80

B4B7BB

9192969B

103104l0B111L72LL71lB119l-20T2T722l-23124t25127128r29131133135136138r39l_42L47150151153156165r67L7TL76180LB4l86187189190191192

L9.915 .919 .614.913.5L3.216 .513.314.313.114 .815 .ll-4.515 .5t2.I14.716.313.011.314 .8L2.814. B

13 .1l-2.413 .113 .0L4.6L2.BL2.TL2.213 .411.518.3It+.5L4.414.611" 316 "714.216 .513. B

15 "7t5.713. B

14.39.9

1r.913 "2

5.L24.9s4 "795 .955 "234.936.726.026.285,79's.676.486.795 "725.25s.9Bs.485.925.585 .334.695.824.825 .005 "625 .686.r75.714.975.6r5.525.785.625.625.725.076.L45 .896 .185 .655.355 .114.985.474.926 .475. B34. B5

13.3 115.6 r14"0 1L4.9 1r3.2 I15.3 1

13.5 113.8 113.9 115.5 113.5 116.1 I16.3 It4.4 I18.5 112.8 I14.8 113.4 1L5.7 112.0 012.8 013.3 016.3 013.0 013.1 013.0 0L6.4 0r7.5 0L3.6 014.1 013.3 0r7.5 016.3 015.3 013.0 013.6 01s.6 014.0 018.1 0L2.6 0r3.3 0r4.7 013.5 013.5 018.5 0L3.6 0

L2.9 0L4.3 0

000000000000000000000000000000000000000000000000

45

TABI,E ] OC A RANKTNG O¡' SELNC TED PLA]:{TS ON A BASIS OF THE

2O OT}'SPRING ]\[EAN YIELD TOCETIIER \^II'I'H THET-R

YIì]LD AND I¡REQUENCY OF Rþ]SEI-ECTION

Sele-ct-.ecl YieldPlantNo.

F.R. 0f f spr:ingMeanYield

Selected Yíeld !-.i{PlantNo.

OffspringMeanYield

6.526.516.506.506.49o .486.476 .476 .476 .466 .426.4r6. 406.396 .386.376. 3s6.3t+6.346.316.306.286,286.286.276.266.256.236.2r6.216.2.r6 .186.L76.r76.L46.146.146.L26.126.r26 .116.106 .086 .086 .06

15.9 318.1 1

11.9 314.3 2'r2.4

2-

15.1 0L2.4 2

20.7 2

9.9 018.4 11s.7 41.0.9 313.8 1L4.2 2

15.8 II2,B 2

1B.B Ir3.7 2

L5.7 I16.3 I16.1 1

12.6 2

18.6 r.

74.3 014.0 311.0 2

14.0 1t4.5 2

L2.7 Ir2.9 119.0 2

r4.2 015.1 1L4.6 013.3 2

13.1 0r1.3 016.6 2

16.5 0Lz.L 2

Lt+.g 2

15.0 1

19.5 2

13.6 Z

12.6 2

13.5 1015.0 6

1.4.s 4

t-4. B s18.5 5

1s.9 6

16.7 3

15.6 6

L9.2 3

14.6 2

13.4 3r4,5 2

L6.9 4L6.7 4

14.8 3L2.4 2

13.0 3

13.C 2

1B.l 2

14.6 2

13.9 3

L1.L 113.6 2

18.3 3

L6.7 2

72.7 I11.9 3L4.5 0L2.9 311.7 3r4.2 3L7.5 014.0 3L6.2 5

14.8 1

15.6 5r3.3 2

14.r 2

14.0 2

L2.3 2

15.3 1r2.3 316.0 IL6.9 316.5 4

11l-54r70r49115

20100L62

45L77

24130743157

996469

L26LB2

269732

L3213

1402966

tL7113

23148

3059

18379

101110

956568

L573649

1683

7 .997 .807 .667.497.387.3r7.rB1 .r37.r27 .067.037 .O26.996.996.996.956.936.9L6.9r6. BB

6.866 .856. B36.826 .816.806 .806.796.746.736.736.7L6.696.696.606.606 .606.596 .586.576.576 "s66.556.536.52

11485T2

15514s].r2

190

190515073

]-60L52

42B33B54

188173t72

6167

104106L46L74

271522

116L77

77135

IB19

165709B

1637.02

25B9

1077

I+6

TABLE lOc Conti.nued

Selected YieldPlantNo.

F.R. OffspríngMeanYield

Sel.ected YieldPl-anÈNo.

F.R. Of f springl'leanYield

13L728

10310

rc47B

L204291

105r22

2

4L67r69

58158191

7594

r21'L66

602IBO

10B.l.47

6

9337

185118ls3r36109133

72111

40r764t

131150151139161

4655

59L23

31rt9

5644

r37r42T2LIB7

B186166335

178180

57124L4t-

5376L4

1r.992

8

7L5

L7574B4

rB4156

5233

r29186l-34138

8796

9189

34t92

39I28

8Bl-25

s .58_s.585.575.575 .5/+5.525.525.525 .485.475 .4-55.45_5 .435.415 .405.385.35s .345 .335 .325.285.28s.275.255.235.205.18s.165.1s5.135.7.25. tl5 .075 .04_s .005 .004. 9B4.974.974.9s4.934.924.924 .874"854.824 "824.794.69

14.0 311.3 013.6 q].4.B 113.0 013.6 014.0 113.4 0L6.3 013.8 0L2.9 114.9 I13.0 0r2.2 2

14.5 2

L2.8 1

13.8 0r4.7 014.8 0I4.9 I18.1 013.6 016.3 0

I2,I O

13.5 013.3 013. -5 0r2.0 018.5 013.5 019.9 0t5.7 0l_4.6 0

14.0 014.r 0r2.4 0

L5.7 015.6 1Lz.L O

15.9 0L3.2 0L2.3 1

r4.3 0r3.3 0l-3.2 016.3 013.l- 019.6 0Lz.B O

6 .046.046.O46.026 .006 .005 .985.985.975 .955.945.92s .90s .895 .895. 885 .83s .835. B35.825.825 "825.825 .81.5 .805 .80s.795.785.775. 755.745.745.725.725.7L5 .705 .685.675.675 .665 .65s.645.625.625.625 .6r5 .605.sB5.58

18. 3

12.7L6 "413. 317 .0L3.2L2.9l-4.l]-4.7L4.912.513.011. 7

11.6t6.713.sL7 .713 .511.918.513.914 .815.sL5.713 .014.313.11t .5L2.B16 .8ú.513.415 .514 .412. B

13. 313.015 .014.813.616.s15 .313.1tB. 3

14. sL2.213"915 .612.6

3I002

1002

01.

0I301110010II00000101000I02

0I000000I00

47

believe conditions t¡oul-d be deÈrímental Ëo seed set. To determine

whether the lack of seed set 1n 1975 rvas due Lo self-incompatibility

or the envíronrnent ln the bags, in 1976 plants Inrere üested for

seed set under bags but wj-lh the possibílity of cross-poll-inati'on'

T\,ro to three Èillers frorn diffeïent plant were enclosed in one bag'

From fifteen tests only five bags provlded seed. The other ten bags

contained emPty spikelets as with the selfed plants '

Thistest'cannotberegardedasconclusíve.Thebagswíthout

seed rnay have been placed on Èhe plants that were mutually incompatible'

or their spikes may have had different blooming periods and cross-

pollination r^ras inadequate. However, there hTas no evidenc'e from these

aÈtempts that self-pollínated seed could be produced'

4B

DISCUSSION

1. Mass sele.ction in I^l . ryegrass

A point that rvill be made in tÌre discussíon ís thaÈ these

results on mass selection within vü. ryegrass indicated a greater

improvement- in yield and therefore a greâter efficíency of selec-tion

than is normal with corn. The reasons for the difference will- be

discussed.

In Èhe literature revíew it was suggested that the íneffective-

ness of mass selection for yíe1d of adapted varietíes ín corn resulted

f ron:

(Í)

(ii)

(iii)1960)

Other suggestions referred to the genetic basís of yield such

that if yield was largely determíned by non-additive effects or over-

dominancerprogress from rnass selection would not be expecte-d

(Gardner 1961). Gardner considered that overdomínance !üas not of

major importance as he found that in open pollinated varieties

relatj-vely la.rge amounts of additive genetic varíance for yield exist"

Four cycles of mass selection resulted in 4.9 per cent gaín per year

over the origínal varietíes.

In the present study of w. ryegrass mass selection for one cycle

resulted in 13.8 per cent gain in yield. Here selection was c.aËr:'.ed

out before anthesis as suggested by Burton, (L974) and it was poss.Lirì'.'

to pollinate with select-e<1 plants. Inferior pollen parents \^7ere

the ir-rabi1íty to ide-ntify superior genotypesfrom Ëheir phenotypic appeararÌce

uncontrolled pollination, so that the selecteciplants were pollinated by both inferior andsuperior pollen parenÈs

an excessive selection intensity leading toreduceci population size, and inbreeding (A1lard,

49

eliminat-ecl . This procedure shor¡l-cl have- clQubled the increase- in the

desirablegenefrecluencybetr^¡eenthegene-rat.ions.I^]itht}reselecl-ion

intensíty used, there vTas no possibj.lity of inbreeding depression'

The 4.9 per cent gaín obtained by Gardner (196f) for corn u/as an

average of four years " To srlpport Lhe suggestion that more rapid

progt:ess can be rnacle iD rnr. ryegl'ass usíng the present techrtitlues it

r¡ou1d be an advantage Èo continue the experiment for some' further

cycles of selection.

If the assumption is made that the response of each genotype to

Ëheenvironmentislinear,thepercentageoftheplantsbe.lov¡arrci

abovethemeaninapopu].ationwillberelativelyconstanteve.nif

there is a large environmental difference between yeaÏS. In Table- 6,

the percentage of the plants belor^r and above the mean of the orj-gina1

population 1975 and 1976 is not const.ant. In other words the ef-.fe-cE

of the environment \^ras to cause a change in the populatioD lnean yield

and a change i-n the proportíon of the plant phenotypes in the populatjon

asv¡ell.tr{henacomparison\'/asmadebeËweentheorigínalandthe

mass selected population in 1976 again a difEerence was obtained' The

percentage of plants above the mean in the mass selecLed popr'rlation

ís greater than in the original population in 1976 an'd tl-re original

population in 1975. An iuterpretatÍorr of this result may be Ëhat,

mass selection has improved the population mean and reduce-d the varj'a-'

bility betv¡een Plants.

A sirnitar ca-lculation \^ras made using the mode as the point of

reference. rn a normal curve, the mode and mean are superirnposed' 0r'r

the other hand in a skewecl distribution thel' deviate' The purpose c1:

Èhe calculaËion was to see rshether there v/as any rnovement of the rnnde

tor¿ards the rnean by measuring the plant percentage below ancl above-- t'tle

50

mode. The percentage, of plarrts bei-or,¡ ttre mode in the selected poPlr-

lation r¡/as great-er than irr tlre unselectecl populatíon (Tab-le 7) ' T-r:

other words, afLeÍ selection the rqode. moved towards Èhe meau t-rf the

populatíon.

Based on the theory of population geneÈics, selection shoui<ì

increase the frequency of deslrabl-e genes and if randon mat-ing is

occurring in the selected population, the-n the form of the clisì:ribut:i orr

should change from a posltive skew toh'ards norlJality as the prrpulaLian

becomes more homogeneous.

IÈ is unlikely that the selected population woul-C become cnn-

pletely hornogeneous and norrnal, because v¡. ryegrass is se'1f-incont¡:atib--l e

and repïoduces by cross-pollination each year. I1- could not be

reduced to a síngle genotype. Furthermore, as pointe'd out by Koyatna

& Kira (f-956), even a totally homogerteous poprrlat:'-on in Ehe genelic

sense may show a skew.

2. Plant assessments

In this study three forms of ranking \^rere consj-dered in eval.u¿itf-j!Ìi

plants and their pïogeny (Tab1e 10). I'r:om Table 10a it c-an be seen

thaË a ranking based on the yíelds of the mass selected plants \'ras nctr'

closely associated with the yíelds of thei.r pTogeny. Qne exp1aÍìêtjroIì

is that some of Èhe selecÈed prla¡ts by chance \,Jere g,ro\,ln in a favour*

able microenvÍronment. Plants tTrat were highiy ranked on a basis of

their yields under mass selection (e.g. plants 90, 84, BB eEc.) trere.

not highly place-d when rankirtg was b¿se-d on. lhe offsprj.ng mean yí-e'lds

(Table 10c). understandably the rankj-ng based on the f::equency cJl r:cr-

selection r¿as relatively consísLent r,rith. the r:ankirrg trf t.hc-' atera¡;e

progeny yield (tabte fOb).

5i"

3. Syn thetic variety in I,/. l'Yegrass

'Itre feasibility of testíng the potential, constituents of a

synthe-tíc variety for their cornbini¡g ability depencls on va::ious

factors. These ínclude-; are the plar-rts self-compatible or i'ncompat-

íble; can they be, vegetatíve.ly proPagatecl or propagated only by

seed. Of the possible combinations of these factors three of the four

readily permit te.sting for combíning abilíty. Thus, if the plants ar:e

self-.compatible they may be selfed and crossed and the selferl seed

may seïve as a means of retainíng the parental genotypes. ft js

advantageous vrhen attenpting to produce synthetic varieties íf the

genotype.s once selecÈed can be retained in the form of int¡red lines

or clones.

The fourÈh combínation, involving incompatibilífy and no vege-

tative- propagation represents the situation í-n w. Tyegrass. coit-

ventional combiní.ng ability tests are difficult to perform as the

plants are self-incompatible and furthe-rmore the life cycle of the

vegetatively propagated plant is not renewed. However, a Progeny

test, giving a measure of combíníng ability was attempted. The-

relevant results are given in Table 9. It is based on the frequency

with which the open-pollinated progenies would have been reselected.

For example, ouÈ of the 20 plant progeníes of the selected plants

Nos. 11, 20,147 and 3, the number of offsprÍng that I'ould have been

reselected was 10, 6, 5 and 4 respectively. By using this progeny

test, plants rvhose offspring having a high resel.ection frequency can

be regarded as plants which have a high general combíning abíli-tv.

Residual seeds from these selected plants could be coutporrncle.d a-nd

release<l as a ne\v syntheti.c variety or they coul.rl be used for the nexÊ

selection cyc1e. Plants whose offspr:ing have a low frequency of

52

reselectiolr uould be regarded as planËr; rvith only a hí-gh specifìc

combining ability. Hor+ever, j-t is difficult to exploit these plants

as hybríd v¿rrieties as the pare-ntal plants cannot be retained as

lines or clones. In addition ru. ryegrass is an annual plant and

combinations, or specific crosses could not be produced re-pe-atedly'

Other factors which influence the assessment of a plant anrl

whi-ch should be considered when making progeny tests are:

The number of progeny that are assessed' fn thi¡'istudy from each mass sele-cted plant tvrenty offspring\^rere gro$7n, but this number could be increased forgreater precisíon.

selection intensity. The intensity of truncatíor1could be varied. Here the truncation was basecl olr

the best Èhree- of the 49 plants ín each subplot'

The size of rhe subplot. In this study a 49 plantplot was used occupying 87.5 x 87.5 cm' Plot sizeÀnould be considererl in relation to soil uníformityand selec.tion ínÈensitY.

4. Alternative mass selection Pro cedure in W. ryegrass

Based on what has been carried out in this studY¡ an alternative

mass selection procedure could be suggested'

The procedure is detailed below (Fig. 10). In the third year

of thís procedure, information could be obËained on addití-vity'

inbreeding depressíon and combining atrility'

5 Purpose and expectaEion from the th i-r:d year f-est-íng (see page 56)

(1) The first synthetic variety (the syn. 1) should have

a higher yield than the unselected popr:1ati.on. IEs con-

stituents will have been further Èested and selecÈed

than the offspring ín the present study which showed

an íncrease ove-r ttre- original populat'íon'

(2) Population 3 should have a higher yi-e'1d than population

1. The reasoning is the same as that gíven fo:: the

53

Syn. 1. FurLher morc if yield of popul'at-ion 3 ís

hígher than the yield of Syn' 1, it ne'ans there hae

beenafurt.herÍncre-asej.nthedesirablegenesor

additive factoïs for yield' If this occurs' then a

secon<l synthetÍc varíety (Syn' 2) could be formed'

If yield of the populaÈion 3 is equal to Syn' 1" it

meanstherehasbeennofurtheracc'umulatíonoftlre

desírable genes. On Ëhe other hand if poprrlation lÌ

is lower than Syn. 1 and population 1, iÈ mear¡s that

inbreeding depression has occurred' The genotypes Ín

population 3 have become homozygous and homogeneous as

a result of the selectíon.

(3) Population 4 should have a lower yield than Syn' 1 and

population 3, and be at leasL the sane as population 1'

The material ín population 4 will have beer' selected

butshouldnotbesouniformastoleadtoirrbreedirrg

depression and a low yielder than population l'

If the yíeld of population 4 is greater than Syn ' I

and population 3, it means there Ís no need to make

assessmerits of combining ability ' Mass selectj on

alone will have been the cause of the improvement'

Further mass selection could be carrÍed out on a basi's

of truncatíon for Yleld.

(4) Population 5 should have a yield lower: than Svn' I

populations 4 and 3. IÈ should be símilar Èo popu-

lation l. It is a population formed from genotypes

which had been an average or l.ow yíelr1 when tesfecl

in the se-cond, year. If r:he yíeld of population 5 is

54

hÍghe.r t-han populations 4, 3, Syn' I and 1, í't means

there is rro differe¡c.e between lorv and high frequency

of rese-lection.

(5) ì,Iith populatíon 6, i.t is possible to checlc whether

ir.rbreeding depr:essiou is occurring with this level

of selection in rv. ryegrass'

All of the information whích could be drav¡n froru the pÏocedure

presented would be informative for further breeding for yield in

\^7. ryegrasS .

i

55

-EIq!BL-LO

First Year

Procedure:

Seed of Original ?oPulation

unselected population

1. Forage evaluation of Plants.

2, Mass selection based on the best n plants out ofan m-p1ant p1ot.

3. 'Move the selected plants to the glasshouse beforeanthesis.

TesÈ for self-inc.ompatÍ.bi1.ity by plac-ì-ng a fewtillers under bags with the objective of findingself fertile lines and the production of inbreds'

As a control to this assessment of fertiliÈy a fewtíllers should be cross-pollinated under bags '

The rest of the tillers should be left open for: inter-crossing.

The seed produced by each pl-ant r^¡ould be kept se-parately.

Seconcl Year

Procedure:

Yield test the unselected population and the selectedpopulation.

Undertake mass selection based on the best n plantout of m plants PIot.

Testíng of c.ombiníng abilíty based on the freclue-rrcyof reselectíon o-f the best n plant out of m plantplot-.

¡,1rt

I

4

5

6

7

1. unselected PoPulation 2 selecte-d progenypopulation

{

i

TI

I

1

2

3

r

56

FIGURF, 10 Con Ëínue

4 Before anthesis in the second year seleeted plantswould be grouped for pollination. The groups wouldbe:

(a) Among plants with high frequency of resel'ectj-on(high I'.R.) .

(b) Among plants with low frequency of reselection(low F.R.) .

(c) Cornbination of (a) and (b).

of each se.lected Plant.(d) Among the. sibs

5 Followe-d by procedure Nos.the first year.

4, 5, 6 and 7 rnentione<l j-n

A first syntheti_c variety could be formed by compoundingremnant seed of plants having a high frequency j'n Lhe

progeny fest.

Third year

In the third year there are síx populations for testing" Fíveof Èhem derived from second year (a to d and the synthetic) arrd <;ne

from the first year (unselected population) '

6

firstsynthetícvariety(Syn. 1)

combinationof low andhigh r'.R.populatíon

sib c::ossfrom eac.hsele-c Eedpopulation

híghF. R.population

1owF. R.populatícn

r¡nselectedpopul.ation

1 2 3 54 6

Populatíons

rI

I

I

Procedure:

1. Yield testing ofPoPulation as a

2. Continuation of

the five populations rvith the or:í-gina1.contro 1.

the procedure used in the second year.

51

REFURENCES

Allarcl, R.W., 1960, PrinciPlesJohn lnlileY & Sons Inc.

of Plant Breeding.

Altard, R.W. & Bradsharv, A.D. 'envi ronmental i-nteractionsCr:op Sci. 4: 503-508.

Allard, R.w., Lg67. Popul.atíon sÈructure. and Perforniance i-n crop

Plant-s.Cíencia E Cultura 19. No.l: 145-150'

Andrus, C.F., 1963. Plant Breeding Systens'Errphytica 12: 205-228.

Anonymous, L973. Princíples of Plant Breedi'ng'course notes by staff of the universities of Adelaide'Melbourne ancl

-sydney at Bïawijaya Uníversity' Malang' Java'

Ind.onesia. Australian Asian universiÈie-s co-operati<¡n sche-me '

Brewbaker, J.L. , L964. Agricultural genetics'Prentice-Hal1, Engelwood Cliffs, N'J

Burton, G.l{., 1970. Breedíng subtropical species for increased animal

production.Prox. XI InsË. Grassland Congress: 456-463'

Burton, G.W., Penny' L.ll ., Flallauer, A'R' and Eberhart' S'A'' 1977'

Evaluatíon of synthetic population deve.loped from maízÞ. varicty(BSK) by two methods of recurrerrt selecËion'Crop Sci . 11: 361-365.

BurËon, G.W., 1g74. Recurrent Restri-cted. Phenotypic Selection (R'R'P'S')increase forage yields of Psenacola tsahíagrass'Crop Scí. I4z 83f-834.

coc.kerham, c.c., 1956. Analysis of Quantítative Gene actíon"Genetics in plant breeding No.9: 53-58'Bookhaven SYnPosia in BiologY '

Comstok, R.E. and Robinson, H.R. , Lg52' Genetic parame-ters' theirestimation and significance'Proc. VI Int. Grassland Congress: 284-29L'

Donald, C.M., 1963. Competition among crop and pasture planls'Advanc. Agron. 15: 1-118.

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the United Nations.

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APPENDIX 3

PLANT i{U" TllE YIELD 1'lí: TTË 2) 1f-tSPRING(SUf4 Ul- 3 HARVf:Sï5} 1976"

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