T h e o r . Ap p l . G e n e t . 50, 35-40 (1977)

9 by Springer-Verlag 1977

Genetic Relationships among Three Chlorophyll-Deficient Mutants in Peanut, Arachis hypogaea L. P . Y . P . Tai, R . O . H a m m o n s a n d R . S . M a t l o c k U n i v e r s i t y of G e o r g i a C o l l e g e of A g r i c u l t u r e , C o a s t a l P l a i n S t at i o n and the ARS, USDA, Tifton, and the O k l a h o m a A g r i c u l t u r e E x p e r i m e n t a l Station, S t i l l w a t e r , O k l a h o m a ( U . S . A . ) S u m m a r y r The g e n e t i c r e l a t i o n s h i p s of t h r e e c h l o r o p h y l l - d e f i c i e n t mutant p e a n u t s , l u t e s c e n s ( l u ) , a u r e u s (a__u_u), and v i r e s e e n t (v) w e r e s t u d i e d u n d e r f i e l d and g r e e n h o u s e c o n d i t i o n s . The F~ p l a n t s f r o m c r o s s e s b e tween t h e s e m u t a n t s p r o d u c e d p h e n o t y p i c a l l y n o r m a l g r e e n . In F~, a u r e u s • v i r e s c e n t s e g r e g a t e d 675 n o r m a l g r e e n : 225 v i r e s c e n t : 45 a u r e u s : 15 v i r e s c e n t a u r e u s : 64 s e e d l i n g l e t h a l , and l u t e s c e n s • v i r e s c e n t s e g r e g a t e d 45 n o r m a l g r e e n : 15 v i r e s c e n t : 3 l u t e s c e n s : 1 s e e d l i n g l e t h a l . ( L u t e s c e n s p e a n u t s w e r e s e e d l i n g l e t h a l in the f i e l d . ) As p r e v i o u s l y r e p o r t e d , the F2 of a u r e u s x l u t e s e e n s g a v e 225 n o r m a l g r e e n : 15 a u r e u s : 15 l u t e s c e n s : 1 s e e d l i n g l e t h a l . The t h r e e c h l o r o p h y l l - d e f i c i e n t f a c t o r s (a._.~u, l u , and v) show independent i n h e r i t a n c e . The r e c e s s i v e c o m b i n a t i o n s f r o m the p a r e n t a l t y p es b e t w e e n a u r e u s and v i r ' ~ s c e n t and b e t w e e n a u r e u s and l u t e s c e n s would p r o d u c e p l a n t s with a c o m b i n a t i o n of t h e i r r e s p e c t i v e p a r e n t a l c h a r a c t e r i s t i c s , but the r e c e s s i v e c o m b i n a t i o n b e t w e e n l u t e s c e n s and v i r e s c e n t was n e a r l y a l b i n o . The v-au and l u - a u s e e d l i n g s h a v e a l o n g e r l i f e span than the z~-lu s e e d l i n g h a s . The g e n o t y p e s f o r the t h r e e m u t a n t s a r e t e n t a t i v e l y i d e n t i fied as l u t e s c e n s W Au~.Au~. Au2Au~ lu:L lu., luz lu2 L~.L:LL~L~ , a u r e u s W a u l a u • au2au~ Lu:~Lu~ Lu~Lu2 LzL~. Z212, and v i r e s c e n t vv Au~.Au• Au2Au~ lu~ lu~. Lu2Lu~ l~l~. L2L2. Key w o r d s : P e a n u t - C h l o r o p h y l l - Mutants

Introduction

Araehis hypogaea ssp. hypogaea and A.hypogaea ssp. fastigiata v a r . vulgaris, l e t h a l s that r a n g e f r o m z y g o t i c

In i n f r a s p e c i f i c h y b r i d i z a t i o n s b e t w e e n

s t a g e to y e l l o w s e e d l i n g a r e u s u a l l y o b s e r v e d in the F 2 g e n e r a t i o n u n d e r f i e l d c o n d i t i o n s . A r e v i e w on c h l o r o p h y l l - d e f i c i e n t i n h e r i t a n c e in t h i s c r o p h as been m a d e by H a m m o n s ( 1 9 7 3 ) . The o b j e c t i v e of this study was to e x a m i n e the i n h e r i t a n c e m o d e l and to d e t e r m i n e the g e n e t i c r e l a t i o n s h i p of t h r e e c h l o r o p h y l l d e f i c i e n t m u t a n t s in peanuts.

M a t e r i a l s and M e t h o d s Three chlorophyll-deficient mutants, virescent, aur e u s , and l u t e s c e n s , and t h e n o r m a l g r e e n k r i a k l e l e a f p e a n u t , w e r e s t u d i e d . V i r e s c e n t , which o r i g i nated from a radiation-induced mutation (W.C. G r e g o r y , p e r s o n a l c o m m u n i c a t i o n ) , s h o w s the y e l l o w i s h - l i g h t - g r e e n in the s e e d l i n g s t a g e and the young l e a f l e t s with s o m e w h a t albino r a c h i s e s and m i d r i b , a s r e p o r t e d by P a t i l and B o r a (1963) and P a t i l ( 1 9 6 9 ) .

Young l e a f l e t s b e c o m e g r e e n e r with a g e , s t a r t i n g f r o m the tip and a r o u n d the m i d r i b s . The y o u n g e s t t h r e e to f o u r l e a v e s c o n t i n u e to be c h l o r o p h y l l - d e f i c i e n t , but t h e l o w e r l e a v e s a p p e a r to be n o r m a l g r e e n . B e n e d i c t and K e t r i n g ( 1 9 7 2 ) r e p o r t e d that c h l o r o p h y l l s y n t h e : s i s in the v i r e s c e n t l e a v e s sh o w s a 7 2 - h o u r l a g p e r i od b e f o r e t h e o n s e t of a p h a s e of r a p i d c h l o r o p h y l l a c c u m u l a t i o n as c o m p a r e d to the n o r m a l g r e e n l e a v e s . The c h a n g e to g r e e n i s h a s t e n e d a s t e m p e r a t u r e i n c r e a s e s and s u n l i g h t i n t e n s i f i e s . A u r e u s , which was found in a plot of PI 268637 (Stone 1968), has young l e a v e s of n o r m a l g r e e n c o l o r that t u r n y e l l o w a s t he l e a f a g e s . L u t e s c e n s 0026 was i s o l a t e d f r o m F~ p r o geny of the c r o s s PI 259662 • k r i n k l e l e a f (Tai and Todd 1972). K r i n k l e l u t e s c e n s was found in a n u r s e r y plot of PI 234422 (Tripp 1968). Both l u t e s c e n s m u t an t s show a p a l e g r e e n l e a f - c o l o r t h r o u g h o u t all s t a g e s of l e a f d e v e l o p m e n t . L u t e s c e n s 0026 ha s s m o o t h l e a f l e t s , w h e r e a s k r i n k l e l u t e s c e n s has c r i n k l e d l e a f l e t s . K r i n k l e l e a f ( H a m m o n s 1964) has c r i n k l e d l e a f l e t s with n o r m a l g r e e n c o l o r . C r o s s e s a m o n g m u t a n t s and with n o r m a l g r e e n w e r e m a d e in the g r e e n - h o u s e . F~ p l a n t s w e r e g r o w n e i t h e r in the g r e e n h o u s e o r in the f i e l d a s n e c e s s a r y to d e v e l o p F~ s e e d s . F o u r F2 p o p u l a t i o n s , two e a c h f r o m a u r e u s • v i r e s c e n t (C 298-2 and C 2 9 8 - 3 ) a n d k r i n k l e • v i r e s c e n t (C 296-2 and C 296-3) w e r e f i e l d - g r o w n ; the o t h e r F~ ' s w e r e g r o w n in the g r e e n h o u s e f o r p h e n o t y p i c c l a s s i f i c a t i o n . To e n s u r e u n i f o r m g e r m i n a t i o n , the F~ s e e d s w e r e t r e a t e d with e t h y l e n e and kept f o r 24 h o u r s in the s e e d g e r m i n a t o r . F~ phenotypic f r e q u e n c i e s w e r e t e s t e d with c h i s q u a r e f o r g o o d n e s s - o f - f i t to p r o p o s e d g e n e t i c r a t i o s .

T h e o r . A p p l . G e n e t . 50 (1977)

36 R e s u l t s and D i s c u s s i o n

Table 1. Fe p h e n o t y p e s and X2 a n a l y s e s f o r l e a f c o l o r in the peanut c r o s s k r i n k l e • a u r e u s

In the v e r y e a r l y s e e d l i n g s t a g e , both l u t e s c e n s and v i r e s c e n t p l a n t s a r e a " w h i t e " phenotype s i m i l a r to

X2

Frequency Population

Green

Aureus

( 15: 1 )

P

C 302-1 C 302-2 Pooled Homogeneity

435 515 950 -

27 30 57 -

0 .1 3 0 0.517 0.597 0.049

.718 .473 .440 .824

a l b i n o , but p i g m e n t d e v e l o p m e n t c h a r a c t e r i z e s t h e i r d i f f e r e n c e s not only f r o m a l b i n o , but a l s o f r o m e a c h o t h e r a s soon a s light i n t e n s i f i e s and t e m p e r a t u r e i n creases.

A u r e u s p l a n t s begin as n o r m a l g r e e n s e e d -

l i n g s . The c o m p l e t e d e v e l o p m e n t of the c h l o r o p h y l l type i s b e s t s e e n in the m a t u r e p l a n t . H o w e v e r , the golden y e l l o w c h a r a c t e r of this mutant type a p p e a r s in the c o t y l e d o n s at the s e e d l i n g s t a g e . The a u r e u s c o t y l e d o n s a r e golden y e l l o w at c r a c k i n g t i m e , but

f o r t h i s r e s e a r c h is a r a d i a t i o n - i n d u c e d m u t a n t . Such

they t u r n g r e e n a f t e r e x p o s u r e to l i g h t . The c h l o r o -

g e n e t i c a b e r r a t i o n has p r e v i o u s l y b e e n d e s c r i b e d by

phyll p i g m e n t p e r s i s t s a p p r o x i m a t e l y two w e ek s and

G u s t a f s s o n (1938) and L o e s c h and H a m m o n s ( 1968) .

then d e c o m p o s e s ,

M o r e o v e r , Hull (1937) found d e f i c i e n c i e s of y e l l o w

l e a v i n g the c o t y l e d o n s y e l l o w

a g a i n . A u r e u s ca n be d i s t i n g u i s h e d u n a m b i g i o u s l y

s e e d l i n s and s u g g e s t e d p o s s i b l e c a u s e s a s unequal

from green as seedlings.

n u m b e r s of e f f e c t i v e g a m e t e s o r d i f f e r e n t i a l v i a b i l i ty of z y g o t e s . The gene s y m b o l v, d e v i s e d by P a t i l (1969, 1973) f o r the g r e e n - v i r e s c e n t r e l a t i o n was ad ap t ed in t h i s study.

I n h e r i t a n c e and G e n e t i c F a c t o r s f o r the T h r e e C h l o r o p h y l l Mutants

B e c a u s e n e i t h e r p a r e n t co u l d p r o d u c e the albino (lethal) seedling per se, each parent presumably c o n t r i b u t e d a l e t h a l f a c t o r . B a s e d upon the p r e s e n t

1. Luteseens. Both l u t e s c e n s 0026 and k r i n k l e l u t e s c e n s h a v e been found g e n e t i c a l l y by Tai et a l . (1970)

r e s u l t s and t h e i r b o t an i cal t y p e s , the l e t h a l s e e d l i n g g e n o t y p e s L i L l l 2 l 2 and llZ1L2L 2 might be a s s i g n e d

to be double r e c e s s i v e to n o r m a l g r e e n . The F 2 p l a n t s

r e s p e c t i v e l y to k r i n k l e and v i r e s c e n t . The c o m p l e t e

from normal green • lutescens segregated for green

g e n o t y p e s m i g h t be vv ZlZ1L2L2 f o r v i r e s c e n t and

and l u t e s c e n s in a 15:1 r a t i o . Both m u t a n t s h a v e an

7V L I L I ~ 2 I 2 f o r k r i n k l e .

i d e n t i c a l genotype f o r the c h l o r o p h y l l - d e v e l o p m e n t factors. Two s y m b o l s , LUllUlLU2ZU2 , h a v e been a s s i g n e d for the g r e e n - l u t e s c e n s r e l a t i o n s h i p .

I n t e r r e l a t i o n B e t w e e n the A u r e u s and V i r e s c e n t Types

2. Antaeus. The a u r e u s • n o r m a l g r e e n ( k r i n k l e ) c r o s s g a v e 15 g r e e n : 1 a u r e u s in F 2 (Table 1) as

It h as b e e n shown that v i r e s c e n t is a s i m p l e r e c e s s i v e

p r e v i o u s l y o b t a i n e d by M a t l o c k et a l . ( p e r s o n a l c o m -

and a u r e u s i s a doubly r e c e s s i v e to n o r m a l g r e e n and

m u n i c a t i o n ) . Two f a c t o r i a l p a i r s , AUlaUlAU2au2, w e r e

F 1 p l a n t s a r e g r e e n . U n d er cool t e m p e r a t u r e and low

p r e v i o u s l y a s s i g n e d (Tai et a l . 1970).

light i n t e n s i t y , the f i r s t two new young l e a v e s on the

3. VireseenV. This c h a r a c t e r is e x p r e s s e d as a 3:1

F i plant show the " v i r e s c e n t " c h a r a c t e r i s t i c s , but

r e c e s s i v e (Table 2 ) . A n o t h e r p h e n o t y p i c c l a s s , albino

t h i s l e s s d i s t i n c t c h a r a c t e r v a n i s h e d a s soon as the

s e e d l i n g , which is l e t h a l , was p r o d u c e d in the F 2 p o p -

l e a v e s b e c a m e o l d e r . In the F 2 g e n e r a t i o n , f i v e d i s -

ul at i o n f r o m c r o s s e s b e t w e e n k r i n k l e and v i r e s c e n t .

t i n ct t y p e s of p l a n t s w e r e r e c o g n i z e d (Table 3 ) . Two

The phenotypic s e g r e g a t i o n r a t i o f o r g r e e n : v i r e s c e n t :

w e r e p a r e n t a l t y p e s , one was l i k e the F 1 h y b r i d , and

l e t h a l is 45: 15: 4, but d i f f e r e n t i a l v i a b i l i t y of the 1ethal

two w e r e e n t i r e l y n ew t y p e s . One was a r e - c o m b i n a -

p h e n o t y p e was p r e s u m a b l y r e s p o n s i b l e f o r t h e v a r i a b l e -

tion of t h e two r e c e s s i v e s ,

n e s s of c h i - s q u a r e f o r d i f f e r e n t p r o g e n i e s f r o m the

o t h e r was l e t h a l . Although v i r e s c e n t - a u r e u s p l a n t s

s a m e c r o s s - c o m b i n a t i o n . The a b e r r a n t s e g r e g a t i o n

r e s e m b l e v i r e s c e n t in t h e e a r l y s e e d l i n g s t a g e , b e -

m a y be a t t r i b u t e d to the fact that t h e v i r e s c e n t u s e d

c a u s e of t h e i r p o o r v i g o r , " a u r e u s " t r a i t b e c o m e s

virescent-aureus;

the

P.Y.P.

Tai e t a l . : C h l o r o p h y l l - D e f i c i e n t

T a b l e 2. F2 p h e n o t y p e s a n d X~ a n a l y s e s

Arachis hypogaea L.

Mutants in Peanut,

in the peanut cross

F requency

37

krinkle x virescent •

X~

Population

Green

Virescent

Lethal

( 45 : 15 : 4 )

P

(3 : 1)*

P

C 296-1 C 296-2r C 296-3 r Pooled Homogeneity

477 240 342 1059

164 92 103 359 -

16 20 14 50

16. 451 1. 5096 8.8574 20.3548 6. 4 6 3 2

.0003 .470 .012 .00001 .049

0 . 117 0.301 0.816 0.076 2. 158

.943 .521 .665 .962 9 707

* Lethal plants excluded Field-grown

T a b l e 3. Fe p h e n o t y p e s a n d X~ a n a l y s e s ratio = 675:225:45:15:64)

for leaf color in the peanut cross

virescent

• aureus

(Expected

Frequency

Population

Green

Virescent

Aureus

C 298-1 C 298-2r C 298-3t Pooled Homogeneity

296 366 394 1056 -

87 96 149 332 -

15 21 18 54 -

Virescent -Aureus

Lethal

X~

P

6 6 12 24

16 29 18 63

6.276 5.363 16.341 5.047 22.933

9 .252 .003 .283 .004

t Field grown

evident

much earlier

other genetic factors,

than would be usual.

D u e to

these plants varied in viabili-

cross

w o u l d g i v e 27 g e n o t y p e s o f l e t h a l i t y ,

ing on the combination of the other three chlorophyll

(V, Au1

t y , r a n g i n g f r o m t h e s e e d l i n g l e t h a l to m a t u r e - p l a n t

factors

under favorable greenhouse

that unequal numbers

ditions,

culture.

all virescent-aureus

As noted previously and virescent,

Under field con-

plants died as seedlings.

for crosses

between krinkle

because neither parent could produce

the albino seedling per s e, both virescent carry

lethal gene(s).

krinkle both carry are Spanish

We assumed

and aureus

that aureus and

the same lethal genes because both

(A.hypvgaea

ssp.

fastigia~a

var.

depend-

and

An2).

Hull (1937)also

of effective gametes

suggested or differ-

ential viability of zygotes could cause the deficiency of y e l l o w s e e d l i n g s .

Furthermore,

variant behavior with virescent previously

the possibility of

has been discussed

in the krinkle • virescent

cross.

act the same way in causing abnormal t h e F 2 of v i r e s c e n t

It m i g h t

segregation

in

• aureus.

'vulga-

ris ' ) type. Based on the results

of our study, the F 1 genotype

Vv AUlaUlAU2au2 L1Z1L212 genies reus,

of green,

virescent,

was proposed. aureus,

F2 pro-

675:225:45:15:64.

W h i l e t h e d a t a s h o w n i n T a b l e 3 do the deviations

ly explained by the deficit of lethals.

Between the Lutescens

and Virescent

virescent-au-

and seedling lethal are expected in the ratio of

not fit this ratio closely,

Interrelation Types

may be large-

Lethality may

In crosses between lutescens and lirescent, the F 1 plants were green, but the virescent (not lutescens) characteristic could be seen in the first one and two young leaves on these plants under favorable condi-

occur anytime from zygotic to seedling stage, but

tions. The F 2 segregated four apparent types --

only the seedling lethals are detected and counted. Ac-

green, virescent, lutescens, and the n e w combina-

cording to the hypothesis, the double recessive fac-

tion seedling lethal. The latter died in the early seed-

tors ZIZI Z2Z2 produces lethality, irrespective of

ling stage, but could be identified. At first, the seed-

other chlorophyll factors present. Theoretically~ the

lings were nearly albino, but low concentration of

Theor.

38

Appl. Genet.

50 ( 1 9 7 7 )

T a b l e 4. F~ p h e n o t y p e s a n d X~ a n a l y s e s f o r l e a f c o l o r i n two p e a n u t c r o s s e s , l u t e s c e n s 0 0 2 6 • v i r e s c e n t (C 3 2 1 ) a n d k r i n k l e l u t e s c e n s • v i r e s c e n t ( C 3 2 2 ) ( E x p e c t e d r a t i o = 4 5 : 1 5 : 3 : 1)

Frequency Population

Green

Virescent

Lutescens

Lethal

C 321-1 C 321-2 Pooled Homogeneity

57 105 162 -

16 37 53

5 7 12

1 6 7

0.878 5.454 3.213 3.119

.831 142 9 360 9 374

C 322-1 C 322-2 Pooled Homogeneity

46 51 97 -

14 22 36

6 5 11

2 4 6

3.613 7.288 8.644 2. 257

.316 .063 .045 9

Pooled total Homogeneity

259 -

89

13

10.015 7 . 218

.018 .614

23

T a b l e 5 9 F2 p h e n o t y p e s a n d X~ a n a l y s e s f o r l e a f c o l o r in t h e p e a n u t c r o s s l u t e s c e n s ( E x p e c t e d r a t i o = 2 2 5 : 15: 15: 1)

Xe

P

9

aureus • krinkle

Frequency Population

Green

Aureus

Lutescens

Lutescens

X2

C 323-1 C 323-2 Pooled Homogeneity

64 82 126

3 6 9

3 5 8

0 1 1

0.965 1.142 0 . 403 1.704

.810 .767 .940 .636

chlorophyll developed in a pattern as does the vires-

Interrelation Between the Lutescens and Aureus Types

(vv) p l a n t 9 D u e to t h e e x t e n s i v e a l b i n i s m , t h e n e w b i p a r e u t a l c o m b i n a t i o n ( vv :lu I lu 1 lu 2 lu 2 ) d i e d b e f o r e e x p r e s s i n g t h e l u t e s c e n s ( lu llUllU 21u2) c h a r -

Extensive genetic investigations

acteristic

b y Tai e t a l .

cent

in the older leaves even when light and

temperature

conditions were very favorable.

t i o n of g e n e s " v " a n d

The ac-

"lu" o n c h l o r o p h y l l d e v e l o p -

merit, as shown in this study, was independent, t h a t of g e n e "v"~ w h i c h r e q u i r e s chlorophyll accumulation

between the lutescens (1970).

on the interrelation

and aureus have been conducted Our additional results

i n T a b l e 5. A s h a s b e e n s h o w n , l u t e s c e n s are controlled by double recessive,

but

loci, symbolized

the lag period in

are shown

and aureus

duplicate unlinked

lUl lUl lU2lu 2 a n d aulaulau2au2 ,

respectively. F I plants from crosses between these

(Benedict and Ketring

mutants are all fully green, both in the seedling and

1972), extends chlorophyll deficiency 9 Therefore,

in the mature growth stages, indicating c o m p l e m e n -

the plants died in the early seedling stage,

tary gene action between Au and Lu9 Four phenotypes

as soon

as food stored in the cotyledons was exhausted. F 2 progenies from both lutescens (C 3 2 1 ) a n d k r i n k l e l u t e s c e n s segregated 3 lutescens

occur in F2, green, aureus, lutescens, and the new

0026 • virescent

• virescent

(C 3 2 2 )

i n t h e r a t i o o f 45 g r e e n : 15 v i r e s c e n t

the expected 225: 15: 15: I phenotypic ratio. The pre:

: 1 lethal with a fairly close fit as shown

i n T a b l e 4. L u t e s c e n s duced a smaller than did the cross

0026 x virescent,

chi-square

however, pro-

value in F 2 populations

of krinkle lutescens

• virescent.

combination lutescens-aureus, and segregation fits

sent results confirm the previous conclusion (Tai et al. 1970). The n e w combination (lUllUllU2lu 2

aulaulau2au 2) of the two parental types produced a plant with lutescens characteristics in early seedling stage, but the chlorophyll was gradually decomposed

P.Y.P.

Tai et a l . : C h l o r o p h y l l - D e f i c i e n t Mutants in P e a n u t , Arachis hypogaea L.

39

Table 6. G e n e t i c c o n s t i t u t i o n of a n o r m a l g r e e n and t h r e e c h l o r o p h y l l d e f i c i e n t mutant t y p e s in p e a n u t s

a c c u m u l a t i o n which v i r e s c e n t h a s , but i t s c h l o r o p h y l l

Chlorophyll types

Chlorophyll factors

When t h e s e two p a r e n t a l t y p e s a r e c o m b i n e d , they

Green (Krinkle)

VV AUlAUlAU2Au2 LUlLUlLU2LU2 L1Lll212

Aureus

VV aulaulau2au 2 LUlLUlLU2Lu 2 L1Lll2I 2

Lutescens (0026 and Krinkl e )

VV AUlAUlAU2Au 2 lUllUllU21U 2 LILIL2L2

Virescent

VV AUlAUlAU2Au 2 LuILUlLU2Lu 2 llliL2L2

p r o d u c e albino t h r o u g h the a c t i o n of g e n e s "Zu" and

as l e a v e s a g e d , p r e s u m e d l y due to the "azz" g e n e s . Even under favorable environments,

a c c u m u l a t i o n cannot i n c r e a s e (Tai and Todd 1972).

"lu-au" p l a n t s

"v", c a u s i n g i m b a l a n c e in both c h l o r o p h y l l f o r m a t i o n and a c c u m u l a t i o n . The v'-au and lu-au s e e d l i n g s have a l o n g e r l i f e span than the v--lu plant h a s .

Acknowledgment The a u t h o r s thank T . A . C o f f e l t , D . W . G o r b e t , and W . W . Hanna f o r m a n u s c r i p t r e v i e w . The work was s u p p o r t e d by t h e f o l l o w i n g : The G e o r g i a C o a s t a l P l a i n S t a t i o n , the O k l a h o m a St at e U n i v e r s i t y , the A g r i c u l t u r a l R e s e a r c h S e r v i c e of the U . S . D e p a r t m e n t of A g r i c u l t u r e and the G e o r g i a A g r i c u l t u r a l C o m m o d i t y Commission for Peanuts.

t u r n golden y e l l o w in c o l o r and die in the l a t e s e e d ling s t a g e .

Literature

The s e v e n r e c e s s i v e f a c t o r s f r o m t h r e e m u t a n t s d i s c u s s e d in this p a p e r i n t e r a c t to g i v e the c h l o r o p h y l l a b n o r m a l i t y . The n o r m a l g r e e n peanut plant m u s t then p o s s e s s the d o m i n a n t a l l e l e s of t h e s e f a c t o r s . H o w e v e r , u n d e s i r a b l e a l l e l e s can a c c u m u l a t e in s e l f p o l l i n a t i n g a m p h i p l o i d s p e c i e s if t h e r e a r e t r a i t s c o n t r o l l e d by d u p l i c a t e f a c t o r s ( A s h r i 1968; Hull 1937). The g e n e t i c c o n s t i t u t i o n of t h e s e m u t a n t s i s p r e s e n t e d in Table 6. The p r e s e n t data i n d i c a t e that both a u r e u s and l u t e s c e n s g i v e c o n s i s t e n t l y b e t t e r f i t s of the t h e o r e t i c a l s e g r e g a t i o n r a t i o s than d o e s v i r e s c e n t . Thus, the v i r e s c e n t mutant m a y c a r r y r e s i d u a l b a c k g r o u n d genotypic changes from chromosome structural dama g e , w h e r e a s the a u r e u s and l u t e s c e n s m u t a n t s would not h a v e s u c h a l t e r a t i o n s . The t h r e e c h l o r o p h y l l - d e f i c i e n t f a c t o r s (au,

lu,

and v) show i n d e p e n d e n t i n h e r i t a n c e . The r e c e s s i v e c o m b i n a t i o n s f r o m the p a r e n t a l t y p e s b e t w e e n

aulaulau2au2 and vv and b e t w e e n aulaulau2au 2 and lu 1 lullu21u 2 would p r o d u c e p l a n t s with c o m b i n a t i o n of t h e i r r e s p e c t i v e p a r e n t a l c h a r a c t e r i s t i c , but c o m bination b e t w e e n ~UllUllu21u 2 and vv was n e a r l y a l b i n o . The a u r e u s i s a m a t u r e - p l a n t type mutant and can o v e r c o m e the l a g p e r i o d in c h l o r o p h y l l a c c u m u l a tion of both the l u t e s c e n s (Tai and Todd 1972) and v i r e s c e n t ( B e n e d i c t and K e t r i n g 1972) t y p e s ; t h e r e f o r e , t h e l u t e s c e n s - a u r e u s (lu-au) and v i r e s c e n t a u r e u s (v-au) p h e n o t y p e s a r e e x p r e s s e d . L u t e s c e n s m a y not h a v e t h e d i s t i n c t la g p e r i o d of c h l o r o p h y l l

A s h r i , A. : M o r p h o l o g y and i n h e r i t a n c e of s t e r i l e b r a c h y t i c d w a r f s in p e a n u t s , Araehis hypogaea. C r o p S c i . 8, 413-415 (1968) Benedict, C.R. ; Ketring, D.L. : Nuclear gene aff e c t i n g g r e e n i n g in v i r e s c e n t peanut l e a v e s . P l a n t P h y s i o l . 49, 972-976 (1972) G u s t a f s s o n , A . : St u d i es on the g e n e t i c b a s i s of c h l o r o p h y l l f o r m a t i o n and the m e c h a n i s m of induced m u t a t i o n . H e r e d i t a s 24, 33-93 (1938) Hammons, R . O . : Krinkle, a dominant leaf m a r k e r in the peanut, Arachis hypogaea L. C r o p S c i . 4, 22-24 (1964) H a m m o n s R . O . : G e n e t i c s of Araehis hypogaea. In: P e a n u t s - C u l t u r e and U s e s , 135-173. S t i l l w a t e r , Oklahoma: A m e r i c a n Peanut Res. Educ. Assn. (1973) H u l l , F . H . : I n h e r i t a n c e of r e s t p e r i o d of s e e d s and c e r t a i n o t h e r c h a r a c t e r s in the p e a n u t . F l o r i d a A g . E x p t . Sta. B u l . 314 (1937) Loesch, P . J . J r . ; Hammons, R.O. : A radiation b r e e d i n g e x p e r i m e n t with p e a n u t s . II. I n h e r i t a n c e of m a c r o - m u t a n t s (NC 4 - 1 8 . 5 k R ) . R a d i a t . B o t . 8, 94-108 (1968) P a t e l , J . S . ; J o h n , C . M . ; S e s h a d r i , C . R . : The i n h e r i t a n c e of c h a r a c t e r s in the groundnut Arachis hypogaea. Ind. A c a d . S c i . P r o c . 3 ( B ) , 214-233

(1936) P a t i l , S . H . : I n h e r i t a n c e of X - r a y i n d u c e d v i r e s c e n t mutant in g r o u n d n u t . Ind. J . G e n e t . P l a n t B r e e d . 29, 387-394 (1969) P a t i l , S . H . : C h l o r i n a , a new c h l o r o p h y l l - d e f i c i e n t c h a r a c t e r in g r o u n d n u t . Ind. J . G e n e t . P l a n t B r e e d . 33, 82-87 (1973) P a t i l , S . H . ; B o r a , K . C . : R a d i a t i o n i n d u ced m u t a t i o n s in g r o u n d n u t . I. C h l o r o p h y l l m u t a t i o n s . Ind. J . G e n e t . P l a n t B r e e d . 23, 47-49 (1963) Stone, E . G . : G e n e t i c , a g r o n o m i c , b o t a n i c a l , p h y s i c a l , c h e m i c a l , and o r g a n o l e p t i c e v a l u a t i o n of p e a n u t s , Araehis hypogaea L. P h . D . t h e s i s , O k l a h o m a S t a t e U n i v e r s i t y , S t i l l w a t e r (1968) Tai, Y . P . ; K i r b y , J . S . ; M a t l o c k , R . S . : C h l o r o p h y l l m u t a t i o n s in p e a n u t s , Arachis hypogaea L. II. G e netic analysis. Agron. A b s t r s . , A m e r . Soc. A g r o n . p 21 (1970)

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T h e o r . A p p l . G e n e t . 50 (1977)

Tai, Y . P . ; Todd, G . W . : C h l o r o p h y l l m u t a t i o n s in p e a n u t s , Araehis hypogaea L. I. P i g m e n t a n a l y s i s . C r o p S c i . 12, 13-15 (1972)

T r i p p , L . D . : G e r m p l a s m e v a l u a t i o n and i n h e r i t a n c e s t u d i e s in p e a n u t s , Arachis hypogaea L. P h . D . t h e s i s , O k l a h o m a S t a t e U n i v e r s i t y , S t i l l w a t e r (1968)

R e c e i v e d F e b r u a r y ~ 5, 1977 C o m m u n i c a t e d by A . G u s t a f s s o n

P . Y . P . Tai D e p a r t m e n t of A g r o n o m y Georgia Coastal Plain Station Tifton, GA 31794 (USA) R.O. Hammons A g r i c u l t u r a l R e s e a r c h S e r v i c e , USDA Tifton, GA 31794 (USA) R.S. Matlock % Oklahoma State University S t i l l w a t e r , OK 74074 (USA)

Genetic relationships among three chlorophyll-deficient mutants in peanut, Arachis hypogaea L.

The genetic relationships of three chlorophyll-deficient mutant peanuts, lutescens (lu), aureus (au), and virescent (v) were studied under field and g...
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