Genetical analysis of the efficiency of VA mycorrhiza with spring wheat

Agriculture, Ecosystems and Environment¢ 29 (1989) 273-280 Elsevier Science Publishers B.V., Amste~dam Printed in Czechoslovakia

Genetical

Analysis

of

the

Efficiency

o f V& N y c o r r h i z a

with

SpringWheat G. G.B. HANSKE

Institute for Tropical and Subtropical Crop Science and ~'unal ~Nime at the University of ~ t t i n ~ , Grisebachstr. 6, D-3400 ~ttingen Abstract

The e f f i c i e n c y of VAH depends not only upon the ~nvironmental c o n d i t i o n s but a l s o on the genotype of the host p l a n t . In a pot experiment with a s t e r i l i z e d , P d e f i c i e n t , P - f i x i n g s o i l 22 High f i e l d i n g V a r i e t i e s ~RYVs) and 22 Landraces and wild forms (LHs) of wheat ( T r i t i c u m s s p . ) were i n o c u l a t e d with the m y c o r r b i z a l fungus ~lomus maniboti$ and compared with non-ino,:ulated c o n t r o l plants.Two different P-fertilizer forms were t e s t e d : s o l u b l e m~nocalcium phosphate (MCP) and iron phosphate (FePO4).VAN showed a higher e f f i c i e n c y under P d e f i c i e n c y (FePO~) with most of the Las than with HYVs. The e f f i c i e n t wild v a r i e t y Triticum aestivum rotundatum and the African HYV 'Huanca' were crossed r e c i p r o c a l l y . Chromosomal h e r e d i t y of the YAH e f f i c i e n c y could be okserved. With N C P - f e r t i l i zation d i f f e r e n c e s between the r e c i p r o c a l F~ i n d i c a t e d a l s o p l a s m a t i c e f f e c t s . Introduction nesearch on the g e n e t i c c o n t r o l

of root-microorganism i n t e r a c t i o n s ,

with m y c o r r b i z a l fungi,

r a r e (Krishna et al. 1985, Jensen 1985, Krishna

is still

especially

and Williams 1987).But the VA m y c o r r h i z a l (VAM) mycelium can form the m a j o r i t y of the s o i l m i c r o b i a l biomass ( a ~ a n p a r t i c u l a r l y well i n f e c t e d in f i e l d l i z e r and p e s t i c i d e

a~plioation

from h a r d l y s o l u b l e , relevant fixati~,

for

plant

soil-borne production

1978) and r o o t s of c u l t i v a t e d p l a n t s are s o i l s ,it~, e i t h e r no or low mineral f e r t i -

( S t e f f a n 1985).

VAN can improve the P uptake

or applied P compounds. This i s p a r t i c u l a r l y on d i f f i c u l t

soils

with

a high

such as the acid l a t : , s o l s with high A1 and Fe contents

the humid t r o p i c s and the l l k a l i n e ,

P fertilizer p r e v a l e n t in

calcareous s o i l s of the s u b t r o p i c s

(~oawad

1979). VAN e f f i c i e n c y seems :o depend r o t only upon environmental c o n d i t i o n s an~ e f f i c i e n c y of the VAM fungus but a l s o on the genotype of the host p l a n t a l . 1979,

Sieverdina

cultivars

with

little

1980). need

This i s of

important

fertilizer.

In

for the

the

breeding

of

work d e s c r i b e d

(Gra, et low-input here

the

i n h e r i t a n c e of VA mycorrhizal e f f i c i e n c y in spring wheat has-been i n v e s t i g a t e d .

274

N a t e r i a l s and Netbods Wheat plants were grown in plastic pots containing 2 kg of soil in a greeruhouso under natural sunlight. 1he soil was o:eposed of a P-deficient and P-fixinq, sandy forest soil, with high contents of M, l~t and Fe, mixed with quartz sand in the proportion of 1:1, and limed with CaCO~to 1~ 5.5 . This IM has proved best for the Vl ~corrhizal bingos used, ~Z~vs run.otis, whLch is a very e f f i c i ~ t strain. Before application of the P f o r ~ the soil substrate m sterilized by heat. Two sources of phosphorus with different solubilities were tested, hardly soluble i r ~ phesphate (Fel~) and easily soluble nmocalcium phosphate (IRP), 30 mg P kg-s dry soil. Basic nutrients, s~ficient to eli~nate all nutrient deficiencies except P, were applied in soluti~ after germination. Nycorrhizal inorulum consisted of strorqly infected root pieces of D~torium o~ratum, a tropical, obligatorily e ~ c o t ~ c herb, which were placed wrier the geminating wheat seed. 7he plants were harvested at ~ilk-ripe stage of the grains. Root length was measured by the gridline intersect math0d (Neeman 1966). The roots were stained (PhiliDs and Havman 1970) and placed on a Petri dish under a binocular:The infection rate was ansessed according to the subjective estimation method used in breedL'q prngrans" for mildew resistance in wheat. This is the only possible w~ to investigate VA nvcerrhizal infection in roots of a large number of plants with such an extensive root s ~ t m . The infection rate represents the proportion of root length infected by Vl m~orrhizal fungus. The total infected root length was calculated by multiplying infection rate with root length. Re~esentative sas~les of pmdered straw and ear material lere ashed in ~SO4-Se (~ch and 14bavad 19"~?). P contents ~ r e measured i~tometrically by the vanedate-yellow method (l(umies 1955). P uptake mars P concentration x dry weight.lhe screening ~ t . m conducted in m r 1985 and the ~..~rison of the P and F, generations in m r 1986. TWnt~-t,o High Yielding Varieties and 22 Landraces or wild form were investigated. Results

The $creenina All

44

wheat

varieties

manibotiB. The e f f e c t fertility

conditions

of

were i n f e c t e d

by

the

the symbiosis v a r i e d

(NCP).

The N C P - f e r t i l i z e d

higher dry weights than F e P O 4 - f e r t i l i z e d p l a n t s .

Vk s y c o r r h i z a l under low plants

fungus

Glomu$

(FePO~) and high P

yielded

3 to

5 times

The high l e v e l of a v a i l a b l e P

in the NCP treatments r a i s e d a l s o P c o n c e n t r a t i o n and P uptake ( T a b . l ) . P D e f i c i e n t Conditions (FeP04) Ear and shoot dry weight were increased by ~. manihotis under P - d e f i c i e n t conditions

(FePO4). The uninproved

varieties

(LRs) showed a higher VA mycorrhizal

e f f i c i e n c y (N/NN in ~). The m a j o r i t y of the LRs ranged in t h e i r VA mycorrhizal e f f i c i e n c y of ear dry weight above 130~o 20-30~ more than the mean of the HYVs ( F i g . l ) . But the very high N/NN r a t i o of some LRs was in many cases c o r r e l a t e d with an extremely poor plant growth without VAn, i n d i c a t i n g a high mycorrhizal dependency. The symbiosis

increased

the ear

m~re than

the

shoot

dry w e i g h t

( F i g . l ) , and s i g n i f i c a n t i n c r e a s e s of the shoot dry weights by YAM could only be

2?5

found i n LRs. The e a r p r o p o r t i o n of the HYVs was h i g h e r than of the LRs, but with VAM the LRs could reach t h e l e v e l of HYVs ( T a b . l ) . HYVs were s h o r t e r than LRs, which produced g r e a t e r r o o t l e n g t h than HYVs on average. HYVs showed l e s s v a r i a t i o n i n r o o t l e n g t h and p l a n t h e i g h t than LRs. There were no d i f f e r e n c e s in t h e i n f e c t i o n r a t e of G. maniboti$ between HYVs and LRs on t h e a v e r a g e , but some p r i m i t i v e wheats were more s t r o n g l y i n f e c t e d (above 70q). B~cause of t h e i r g r e a t e r r o o t l e n g t h s t h e LRs showed g r e a t e r i n f e c t e d root l e n g t h . There was no

c o r r e l a t i o n s between VA m y c o r r h i z a l e f f i c i e n c y in terms of

shoot d r y weight and e i t h e r

infection rate

or t o t a l

infected root

l e n g t h . VA~

i n c r e a s e d the P c o n c e n t r a t i o n and P uptake in e a r s and shoot~ in gYVs as well as in LRs. Because of

the g r e a t e r

yield increase

in LRs t h e i r

ear

and shoot P

uptake exceeded t h a t of the HYVs ( T a b . l ) . High P F e r t i l i t y

C o n d i t i o n s (NCP)

The VAM fungus Glomu$ maniboti$ i n f e c t e d wheat r o o t s well fed with P due to NCP f e r t i l i z a t i o n , especially

p o o r l y , when p l a n t s were

and VAN showed p a r a s i t i c

effects,

( T a b . l ) . Eighteen HYVs and s i x LRs showed s i g n i f i c a n t p a r a s i t i s m in

r e s p e c t of shoot dry weight by YAM. I n h e r i t a n c e of

the VA M v c o r r h i z a l E f f i c i e n c y i n a R e c i n r o c a l Cross between an

E f f i c i e n t and N o n - E f f i c i e n t V a r i e t y ~ r l t i c u m aestivum rotUndatum (ro)

(a V A ~ - e f f i c i e n t v a r i e t y }

and 'Huanca'

(Hu)

( n o n - e f f i c i e n t ) were r e c i p r o c a l l y c r o s s e d to o b t a i n seeds f o r the comparison of the P and F: g e n e r a t i o n s . In t h i s and shoot d r y m a t t e r s latter

c a s e in c o n t r a s t

both p a r e n t a l v a r i e t i e s

experiment ~lomus manihotis i n c r e a s e d the e a r

with FePO4 f e r t i l i z a t i o n

as

w e l l as with HCP, in the

t o 1985 where the same fungus a c t e d p a r a s i t i c a l l y

in

(Fig.2).

The w i l d form T. as. rotundatum (ro) y i e l d e d i e s s without YAM than t h e A f r i c a n HYV 'Huanca'(Hu) and both F t s . But ro showed a ~trong p o s i t i v e y i e l d r e s p o n s e to the

symbiosis,

resulting

in

the

highest

~!AM e f f i c i e n c y

for

all

factor

c o m b i n a t i o n s . In the FePO4-treatment the m y c o r r h i z a l e f f i c i e n c y , e s p e c i a l l y f o r

2?6

ear dry weight, was much greater with ro than with Hu and could be transf~rr~:]

by r e c i p r o c a l crossing ~o the n o n - e f f i c i e n t HYV. The e f f i c i e n c y of the F~ rang,.4 between t h a t of the p a r e n t s ,

but the a b s o l u t e dry weight values s u r p a s s e d those

of both parents (Fig.2-Fe?O4) The effect of the inoculation .as different in the MCP treatments. Here only in the case of Hu x ro the M/~M in ~ values of ear and shoot dry weight lay between the low efficiency of Hu and the high of ro (Fig.2-MCP). The non-inoculated ro x Hu - plants

grew very

well,

with

the consequence

that

increase by VAN was low. The better shoot growth of r o x

the

relative

yield

Hu corresponded .it~

the better root growth and higher P-uptake of this hybrid in the uninoculated FePO4

treatment. The significant

differences

in VAM efficiency

indicate that

both chromosomal and pl&smatic genes are involved in the inheritance of the VAM efficiency in .heat (Tab.l).

Differences betveen 22 High Yielding Varieties (IWVs) and 23 landraces (l/bs) of spring wheat in several parmeters, ~ u ~ l a t e d ~ and inocolated ~) ~ t h the Vk s ~ r b i z a l f~gus ~/cms mm~ot~s and fertilized ~th FePO~ or m0oocalcium pb~phate (NCP). * si~LfJJmnt differe~es bergen ]m and Mfor LSD 5~ Ear dr~ weight (mg/pot) FePOq MCP ~m M ax x HYVs LRs LSD 5q LSD 5~

Shoot dry weight (eg/pot) FePO4 NCP NM M NM M

384 ~31 2040* 1307 348 504* 1769 1561 95

205

n

1026

1064

1257" 3533* 3063

3431. 2324

FePO4 KCP M NM M

37 34

377

132

40 60* 56 40* 50 52

3,4

5 4

295

Shoot P - c o n c e n t r a t i o n (mg P/g dry weight) FePO4 MOP NH

LSD 5~ LSD 5~1

(~)

1017

159

HYVs LRs

Ear proportion

3,3 3,3

M

NH

H

4,4" 8,0 "8,0 4,9" 7,8 8,3 0,5

n.s. 0,7

Shoot P uptake (mg P/ pot) FePO4 MCP NM

M

3,4 3,4

NH

M

4,7" 26,8* 18,a 6,2" 26,9 24,9

1,0

3,0 2,0

Z??

~/NM in X

300

e

a

r

s

300,

250i

250

200

200

150

150,

1 O0 .

100!

s

h

o

o

t

50

50

L.ondroces --

~.VA

High Ykddln 9 Vor;4tles

myoorrhizal efficiency in (~r and shoot yield (H/I~ in %) of 22 lligh Yielding Varieties 22 la~raees or Wild ferns of s l r ~ ~heat, ~ a ~ - ~ a t e d (~) or inoculated (14) ~ I ~ ~o~us ~ t ~ s and f e r t i l i z e d w i t h F e ~ . - - - : ~00 % no e f f e c t by V ~

•

(g/pot) O.S]

FePO.

0.2

ILS0 5Z I ~

[300

<

o.

_

m

Y.

5j

:<

= =o ,.

x

p<

(~)

300

150 100

2

100

0

-= o

lehoets

I LSO 5= I

200

50

-

MCP

:1.,\/I

200

•

O-

(g/poilu,=

"~ "~ isheots v,J

ears

150 50

.

'

-

r~ NM

b< I<

~< ~<

n.

-Z..~

~.~

. n . M/NM In IIi

F i g . 2 . Yield and Vk ~ r r h i z a l

e f f i c i e n c y (M/I~ in %) f o r e a r s and shoots of the P and FI genera-

tions of a ~eciprocal cross betwen the African High Yield£ng wheat Variety 'Euanca' (I~) and the v i l d formTriticum aesti~n rotuMatum (re), non-inoculated or inootlate(1 with the t/1 ~eorrtizal fungus Gl~u& ~ o t i $ arid fertilized with Fed~ or NCP. - - - : 100 % no e f f e c t by V ~

Z?8

Tab.2. Effect of ~xx:ulati~ with Glomus manihotis ot~ pa~'~tal and Fs geaerations of reciprocal "cx~ses l ~ t ~ . n the /~rican High Yielding wheat Variety 'Huanca'(HU) and the wild form Tt'iticum ae~tima n~tundatum f ~ t i l i ~ d with I~P, I~ = n~-hx:~'~ated, H = :inoculated with ¢J_~.,= m m ~ t i s * = significant d i f f ~ between NH and H f~r laD ~k o = significant d i f f e r , races betwe~l the reciprocal ¥, for laD St Dry w e i g h t Straw

Ear

H/NH M

'Ruanca'

1471

2705*

184

976

HU x r o r o x HU

962 1610

2418" 2531"

251 157

755 *1158

801

2838*

357

741

LSD 5 q , s e p e r a t e f o r NH and H LSD 5q

508

Shoot

H/NH

NM

r, ae.rotundatum

(mglpot)

in ~

I~

M

Hl(

M

212

2447

4772*

195

1519". 201 1439 124

1716 s2768

3937* 3970

229 143

1542

4157"

270

682

n.s

2067*

1993'

216

n.s. 663

H/~

in ~

269

495 361

Shoot P concentration (ug.Plg) NM )[

983 Shoot P uptake

NM

M

Root length (u/pot) Nil N

'Huan¢~'

2,33

2,51

5,73

11,86"

31

82*

HU x r o r o x HU

1,99 1,96

1,72 1,74

3,48 5,28

6,65* 6,84

26 e37

91" 84*

T.ae.rotundatum

2,01

1,67

3,09

6,86*

32

99*

n.s.

0,3~ 0.41

1,61

bSD 5~,seperate f o r I~H and H LSD 5~

1,72 1,58

n.8.n.8

21

in

279 Discussion The e x t e r n a l hyphae can absorb n u t r i e n t s ,

e s p e c i a l l y with low m o b i l i t y in s o i l

s o l u t i o n l i k e P, Zn and Cu, from a much l a r g e r s o i l volume than the absorbing zone

surrounding

monocotylellous

a

non-mycorrhizal

wheat

with

a

fine

root

{Howeler

branched,

et

extensive

~1.

root

1987).

system

The

has

an

e x c e l l e n t c a p a c i t y for n u t r i e n t uptake from the s o i l and is l e s s dependent upon v e s i c u l a r a r b u s c u l a r mycorrhiza (VAN) than many o t h e r p l a n t s and Khan 1977,

Jacobsen and Nielsen

1983).

Yet t h i s

(Nc Key 1966, Sail

study shows

that

wheat

y i e l d s can be c o n s i d e r a b l y increased by VA mycorrhizal i n f e c t i o n . The FePO4 treatment

used here r e p r e s e n t s

an extreme P d e f i c i e n c y so that

wheat p l a n t s remained small and did not t i l l e r .

the

Although VA mycorrhiza i s most

e f f e c t i v e and a c c e l e r a t e s plant growth, when the P supply in the s o i l is limited as i t

is

here.

Tillering

mycorrhizal e f f i c i e n c y

is

is

an important

estimated

y i e l d component in

by i t s

effect

wheat

and the

on the y i e l d of

the host

p l a n t . Therefore more s c r e e n i n g s under moderate P d e f i c i e n c y are neces~aryo Some wild wheats e v i d e n t l y posses high VA mycorrhizal e f f i c i e n c i e s with b e t t e r grain y i e l d s of the i n o c u l a t e d p l a n t s , and t h i s p r o p e r t y can be t r a n s f e r r e d to a n o n - e f f i c i e n t HYV. But the conclusion about the i n h e r i t a n c e of the VAN must be handled c a r e f u l l y , of

fungus,

plant

fertilization

because t h e r e and

are strong i n t e r a c t i o n s between the genotypes

environmental

conditions.

For

example

with

HCP

G. manihoti$ acted p a r a s i t i c a l l y in the year 1985, but enhanced

growth of both p a r e n t a l v a r i e t i e s

in 1976 with the s u e

experimental design but

d i f f e r e n t weather c o n d i t i o n s . The wide v a r i a t i o n

GZomu$ manibotis. influenced

by

in But

infection the

environment

rate

indicated

susceptibility and

fungus

used

s y c o r r h i z a l e f f i c i e n c i e s of some v a r i e t i e s tested

over s e v e r a l years

to measure

of

a varied

a variety (Nosse

susceptibility

seems to

1975).

for

be s t r o n g l y

Therefore

the

VA

, and P and Fs g e n e r a t i o n s is being

the g e n e t i c a l

v a r i a n c e s . The r e s u l t s are not yet published.

and environment-dependent

280

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/~

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analysis wheat.

of the efficiency of ~grlc. Ecosystems