Grain protein and grain yield as functions of dry matter, plant protein, and chlorophyll characteristics in
elite international winter wheats
by Mohamed Ali Al-Khawlani
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science i n
Agronomy
Montana State University
© Copyright by Mohamed Ali Al-Khawlani (1986)
Abstract:
Wheat grain protein content is important for product utilization and nutritional qualities. Improving
protein content is difficult because of the inverse relationship with grain yield. The relationship of dry
matter, plant protein and chlorophyll characteristics to grain pfotein and grain yield was studied in 66
international winter wheats (Triticum aestivum L.). Field experiments include low, medium, and high
N fertility regimes in 1984 and 1985.
The cultIvars differed for all measured traits. A significant but low negative correlation between grain
yield and percent grain protein was found only in 1984. This suggested simultaneous increases in grain
yield and grain protein could be achieved by selection. Total plant protein was positively correlated
with biological yield and grain yield, but not correlated with percent grain protein.
Nitrogen harvest index (NHI) decreased with increasing soil N levels. High grain protein cultivars were
more efficient than medium and low grain protein cultivars at any soil N level above 110 kg/ha. A
positive correlation between nitrogen harvest index (NHl) and percent grain protein was found in 1985.
Chlorophyll concentration during grain filling period was correlated with grain yield and total plant
protein. High grain protein cultivars had longer chlorophyll duration after anthesis than low protein
cultivars.
Nitrogen harvest index, total plant protein, and chlorophyll duration after anthesis accounted for 88%
and 94% in 1984, and 94% and 90% in 1985, of the total variation among cultivars in grain yield and
percent grain protein, respectively. Simultaneous increases in grain yield and percent grain protein
could be achieved by selecting for high total plant protein, high N-translocation efficiency, and longer
chlorophyll duration after anthesis in wheat. GRAIN PROTEIN AND GRAIN YIELD AS FUNCTIONS OF DRY MATTER,
PLANT PROTEIN, AND CHLOROPHYLL CHARACTERISTICS
IN ELITE INTERNATIONAL WINTER WHEATS
by
Mohamed A li Al -Khawlani
A thesis submitted in p a r t i a l f u l f i l l m e n t
of the requirements f o r the degree
Of
Master o f Science
in
Agronomy
MONTANA STATE UNIVERSITY
Bozeman, Montana
May 1986
Vf
fit
tUsC . 2—
APPROVAL
of a thesis submitted by
Mohamed Ali Al -Khawlani
This thesis has been read by each member of the autho r's graduate
committee and has been found to be s a t i s f a c t o r y regarding con ten t,
English usage, for ma t, c i t a t i o n s , bi b li o g r a p h ic s t y l e , and consistency,
and is ready fo r submission to the College of Graduate Studies.
^
Date
. (U L L < ? -r^
Chairperson, Graduate Cqtipmittee
Approved f o r the Major Department
Date
/ f
^
Head, Major Department
Approved fo r the College of Graduate Studies
Date
-
Graduate Dean
iii
STATEMENT OF PERMISSION TO USE
In presenting t h i s thesis in p a r t i a l
f u l f i l l m e n t o f the requ ire­
ments f o r a master's degree at Montana State U n iv e r s it y ,
I agree that
the Library shall make i t a v a i l a b l e to borrowers under rules of the
Li b r a r y .
B r i e f quotations from t h is paper are allowable without special
permission, provided th at accurate acknowledgement of source is made.
Permission f o r extensive quotation from or reproduction of th is
paper may be granted by my major professor, or in his absence, by the
Dean of L i b r a r ie s when, in the opinion o f e i t h e r , the proposed use of
the material
material
is f o r sch ola rly purposes.
in t h is paper f o r f in a n c i a l gain shall not be allowed without
my w r i t t e n permission.
Signature
Date
Any copying or use of the
M n *
p .f).
iv
ACKNOWLEDGMENTS
I wish to express my sincere appreciation to the fo llowing:
Dr. G. Allan Ta y lo r, my major ad vi so r, fo r his advice, f r ie nd sh ip ,
and encouragement through the course of t h is study.
Dr. Jarvis H. Brown, Dr. C. F. McGuire, and Dr. John M. Martin fo r
sharing t h e i r time, t h e i r advice, and t h e i r enthusiasm whil e serving on
my committee.
The World Bank and Food and A g ri c u l t u r e Organization f o r f in a n c i a l
support that made t h i s program successful.
My w i f e , F a t h i a , and son, Nabel, f o r t h e i r patience and love.
V
TABLE OF CONTENTS
Page
APPROVAL.
........................................................................................................................
Ii
STATEMENT OF PERMISSION TO USE...............................................................................
iii
ACKNOWLEDGMENTS..........................................................................
iv
TABLE OF CONTENTS...........................................................................................................
v
LIST OF TABLES................................................................................................... ..........
vi I
LIST OF FIGURES...............
Ix
ABSTRACT...............................................................................................
x
INTRODUCTION.......................................................................................
I
LITERATURE REVIEW..................................................
2
Nitrogen and Dry M a t t e r ............................
Chlorophyll Content.....................................
MATERIALS AND METHODS................................................................................
C u l t i v a r s ......................................................................................................................
Experimental Design.................................................................................................
Soil N i t r o g e n ............................................................................
Chlorophyll E s t i m a ti o n ..........................................................................................
. Measurements......... ......................................................................................................
S t a t i s t i c a l Analyses..............................................................................................
2
4
6
6
7
7
8
8
9
RESULTS AND DISCUSSION.......................... .....................................................................
11
C u l t i v a r Va ri a t io n in Grain Protein and Other T r a i t s ........................
Relationships Among T r a i t s . . . . .........................................................................
Dry M a t t e r ..............................................................................................................
Plant P r o t e i n .......................................................................................................
Chlorophyll Content...........................................................; ............................
Grain Yi el d and Grain Protein as Function of
Several T r a i t s .......................................
11
14
14
17
20
SUMMARY
25
29
vi
TABLE OF CONTENTS--Continued
:
Page
LITERATURE CITED................................... ............................ .............................................
30
APPENDIX.................................................... ..........................................................................
34
vii
LIST OF TABLES
Tables
1.
2.
3•
4.
5.
6.
7.
8.
9.
Page
Names and o r i g in s of cul t i v a r s used.......................................................
Co rrelation c o e f f i c i e n t s
6
(r) among t r a i t means of
66 cul t i v a r s over 3 soil N levels in two years...............................
)4
Correlation c o e f f i c i e n t s among t r a i t s of 66 geno­
type means over 3 soil N levels in two ye ar s ...................................
21
M u l t i p l e regression analysis expressing percent
grain protein and NHI as function of several
t r a i t s ( 19 84 )..........................................................................................................
25
M u l t i p l e regression analysis expressing percent
grain protein and NHI as functions of several
t r a i t s ( 1985) ..........................................................................................................
26
M u l t i p l e regression analysis expressing percent
grain protein and grain y i e l d as functions of
several t r a i t s f o r 66 cul t i v a r s ...............................................................
27
Mean squares of grain y i e l d and percent grain
protein of three height groups of c u l t i v a r s
combined over years and soi l nitrogen l e v e l s .......................................
35
Mean squares of biolo gic al y i e l d and t o ta l plant
protein of three height groups o f c u l t i v a r s
combined over years and soil nitrogen l e v e l s .......................................
36
Mean squares of chlorophyll concentration (mg/g FW)
at anthesis and e a r l y dough of three height groups
of c u l t i v a r s combined over years and soil nitrogen
l e v e l s ....................................................................................
10.
Mean squares of chlorophyll concentration at hard
dough and chlorophyll duration of height groups
of c u l t i v a r s combined over years and soi l nitrogen
l e v e l s ...................................................................................
11.
Eight t r a i t means of 22 winter wheat c u l t iv a r s
( t a l l group) grown at 3 s oi l N le vel s in 1984.................................
39
vi i i
LIST OF TABLES--Continued
Tables
Page
12.
Eight t r a i t means of 22 wint er wheat c u l t i v a r s
(medium height group) grown a t 3 soi l N levels
in
1984.....................................................................................
13.
Eight t r a i t means of 22 wint er wheat c u l t i v a r s
(short group) grown at 3 soil N levels in 1984......... .....................
41
Eight t r a i t means of 22 win te r wheat c u l t i v a r s
( t a l l group) grown at 3 soil N levels in 1 9 8 5 . . .............................
42
14.
15.
Eight t r a i t means of 22 win te r wheat c u l t i v a r s
(medium height group) grown at 3 soil N levels
in
1985..........................................................................
16.
Eight t r a i t means of 22 winter wheat c u l t i v a r s
(short group) grown at 3 soil N levels in 1985...............................
17.
18.
19.
T r a i t means of 22 t a l l c u l t i v a r s grown in 1984
and 1985 at 6 soil N l e v e l s ...........................................................................
44
45
T r a i t means of 22 medium height c u l t i v a r s grown
in 1984 and 1985 at 6 soil N l e v e l s .......................................................
46
T r a i t means of 22 short c u l t i v a r s grown in 1984
and 1985 at 6 soil N l e v e l s ..........................................
47
i X
LIST OF FIGURES
Figures
1.
2.
3.
4.
5-
6.
7-
8.
Page
Frequency d i s t r i b u t i o n o f percent grain protein
f o r 66 c u l t i v a r s at 3 soil N le vel s ( 1 9 8 4 ) ........................................
12
Frequency d i s t r i b u t i o n of percent grain protein
f o r 66 c u l t i v a r s at 3 soil N lev el s (1 985 ) ......... ..............................
12
The r e l a ti o n s h ip between grain y i e l d and grain
protein (%) f o r high and low grain protein
c u l t i v a r s over 3 soil N levels ( 1 9 8 4 ) .....................................................
15
Grain protein (%) and harvest index f o r high and
low grain protein c u l t i v a r s over 3 soil N levels
( 1 9 8 4 ) ......................................................................................................................
16
N harvest index and grain protein (%) f o r high
and low grain protein c u l t i v a r s over 3 soi l N
le vel s ( 1985) .......................................................................................................
18
N harvest index f o r 3 grain protein wheat groups
at 6 levels of
soil N....................................................................................
19
Chlorophyll duration and grain protein (%) for
high and low protein c u l t i v a r s over 3 soil N
levels ( 1 9 8 4 ) .......................................................................................................
24
Chlorophyll duration f o r high and low grain
protein groups at 3 soi l N levels in two years ...............................
24
ABSTRACT
Wheat grain protein content is important fo r product u t i l i z a t i o n
and n u t r i t i o n a l q u a l i t i e s .
Improving protein content is d i f f i c u l t
because of the inverse r e l a ti o n s h ip with grain y i e l d .
The r e l a ti o n s h ip
of dry m a t t e r , plant protein and chlorophyll c h a r a c t e r i s t i c s to grain
pt ot ei n And grain y i e l d was studied in 66 i n te rn at io na l win te r wheats^
(T rit ic um aestivum L . ) .
Fi el d experiments include low, medium, and high
N f e r t i l i t y regimes in 1984 and 1985.
The c u l t i v a r s d i f f e r e d f o r a l l measured t r a i t s .
A s i g n i f i c a n t but
low negative c o r r e l a t i o n between grain y i e l d and percent grain protein
was found only in 1984. This suggested simultaneous increases in grain
y i e l d and grain protein could be achieved by se l e c t i o n .
Total plant
protein was p o s i t i v e l y corr el at ed with bio lo gi c al y i e l d and grain y i e l d ,
but not cor re la te d with percent grain p ro te in .
Nitrogen harvest index (NHl) decreased with increasing soil N^
levels.
High grain protein c u l t i v a r s were more e f f i c i e n t than medium
and low grain protein c u l t i v a r s at any s oi l N level above HO kg/ha.
A
p o s it iv e c o r r e l a t i o n between nitrogen harvest index (NHl) and percent
grain protein was found in 1985Chlorophyll concentration during grain f i l l i n g period was corre­
lated with grain y i e l d and t o t a l plant pr o t e i n .
High grain protein
c u l t i v a r s had longer chlorophyll duration a f t e r anthesis than low
protein c u l t i v a r s .
Nitrogen harvest index, t o t a l pla nt p r o t e i n , and chlorophyll
duration a f t e r anthesis accounted f o r 88% and 94% in 1984, and 94% and
90% in 1985, of the t o ta l v a r i a t i o n among c u l t i v a r s in grain y i e l d and
percent grain p r o t e i n , re sp e c t iv e ly .
Simultaneous increases in grain
y i e l d and percent grain protein could be achieved by sel ect in g fo r high
t o t a l plant p r o t e i n , high N-t ran sl oca tio n e f f i c i e n c y , and longer chloro­
phyll duration a f t e r anthesis in wheat.
I
INTRODUCTION
Wheat grains c o n s t i t u t e the stabl e food of a large proportion of
the world population.
Therefore, wheat protein represents a major
source of protein fo r both humans ahd animals.
important f a c t o r fo r both baking and n u t r i t i o n a l
wheat.
Grain protein is an
properties of bread
Increasing grain y i e l d and grain protein simultaneously is the
ul ti m a t e goal fo r many wheat breeding programs around the world.
The
simultaneous improvement of grain y i e l d and grain protein is d i f f i c u l t
because of t h e i r inverse r e l a t i o n s h i p .
However, c u l t i v a r s with high
grain y i e l d and high percent grain protein have been obtained (Johnson
et a l . , 1967).
This suggests improving both grain y i e l d and protein
content by select ion
is possible.
Increased grain y i e l d and percent
grain protein may be associated with increased use of nitrogenous f e r t i
l i z e r s or increased e f f i c i e n c y in t ra nsl oca tin g nitrogenous compounds
from ve g eta tiv e parts of the plant to the gr ai n.
nitrogenous f e r t i l i z e r s
Higher costs of
in recent years have drawn a t t e n t i o n to the
creation of genotypes with an improved e f f i c i e n c y of nitrogen
u tilization
(Austin et a l . ,
1977).
This study was conducted to examine the genetic v a r i a t i o n and the
r el at io ns hi ps of characters related to grain y i e l d and percent grain
protein in e l i t e
inte rn at io na l win te r wheats.
2
LITERATURE REVIEW
Nitrogen and Dry Matter
Numerous in vestigators have reported s i g n i f i c a n t inverse r e l a t i o n ­
ships between grain y i e l d and percent grain protein in spring and wint er
wheats (Terman et a l . ,
Busch, 1982).
1969; H a ll o r a n , 1981; B h at ia , 1975; L o f f l e r and
The range o f c o r r e l a t i o n c o e f f i c i e n t s from - 0 . 4 8 to -0 .5 8
suggests no genetic l i m i t a t i o n s f o r improving both grain y i e l d and grain
protein percentage in wheat.
Stuber et al . ( 1962) reported high-
y i e l d in g lines with high grain protein content were found in an F2 poput ion of a high x low protein wheat cross.
This suggests simultaneous
improvement of grain y i e l d and grain protein is possible.
Wheat plants are known to accumulate most of t h e i r nitrogen in the
v eg et a tiv e parts p r i o r to anthesis.
developing grain a f t e r anthesis.
The nitrogen is translocated to the
McNeal e t a l .
(1968) found the
nitrogen content of ve g et a tiv e parts of seven spring wheat genotypes
(leaves, stems, and head ch af f) decreased a f t e r anthesis, while grain
nitrogen content increased.
This is i n d i r e c t evidence f o r translocation
of nitrogenous compounds from ve g et a tiv e organs to the developing
grains.
Translocation of labeled amino acids from culms to grains is
d i r e c t evidence of mass tran sl oca tio n o f nitrogen to the grain (Mikesell
et a l . ,
1971).
Nitrogen t ra nsl oca tio n e f f i c i e n c y represents the a b i l i t y
of 3 genotype to t ra ns lo ca te nitrogenous compounds from the vege ta tive
parts to the gr ains .
The e f f i c i e n c y of p a r t i t i o n i n g of nitrogen between
3
straw and grains is expressed as nitrogen p a r t i t i o n i n g e f f i c i e n c y
( L o f f I e r and Busch, 1982), nitrogen tran sl oca tio n e f f i c i e n c y ( H a l loran
and Lee,
1979 ), or nitrogen harvest index (Desai and Bhatia,
1978).
It
is calculated as the r a t i o of grain nitrogen to t o ta l plant ni tro g en S i g n i f i c a n t d iff ere nc es among wheat c u l t i v a r s were found fo r nitrogen
tran sl oca tio n e f f i c i e n c y (Halloran and Lee, 1979; Dubois and F o s s a t i ,
1981; L o f f l e r and Busch, 1982; L o f f l e r et a l . , 1985)•
Halloran (1981)
found nitrogen tran sl oca tio n e f f i c i e n c y decreased with increasing
a v a i l a b i l i t y of soil nitr og en , but the high grain protein c u l t i v a r s
remained highly e f f i c i e n t at high soil nitrogen le v e l s .
L o f f l e r and
Busch (1982) reported select ion f o r high nitrogen harvest index s i g n i f i ­
ca nt ly improved grain y i e l d in the progeny o f three crosses of hard
spring wheat genotypes.
Grain protein content s i g n i f i c a n t l y increased
in one po pu lat io n, with no reductions in the others.
Nitrogen t r a n s l o ­
cation e f f i c i e n c y can be an important c r i t e r i o n fo r improving grain
yield
(Dubois and Fossati , 1981) , or grain protein content (Halloran and
Lee, 1979; L o f f l e r et a l . ,
1985).
Total plant nitrogen can be considered as an ind ic at or of the
e f f i c i e n c y of nitrogen uptake (Desai and Bhatia,
1978).
The phys iolog i­
cal basis f o r high grain y i e l d and high percent grain prot ein in wheat
appeared to be associated with gr eat er nitrogen uptake.
Increased to ta l
plant nitrogen at maturity or more e f f i c i e n t and complete translocation
o f nitrogenous compounds from the v e g et a ti ve plant parts to the grains
are l i k e l y .
Total plant nitrogen at m a tu r ity was p o s i t i v e l y correl ate d
with grain y i e l d , but not s i g n i f i c a n t l y corr el at ed with grain protein
content (Desai and Bhatia, 1978; Cox et a l . ,
1985b; L o f f l e r et a l . ,
4
1985) •
This suggests select ion f or high t o t a l
plant nitrogen could
improve grain y i e l d without reducing percent grain p ro te in .
al.
Johnson et
(1967) found nitrogen uptake and nitrogen tran sl oca tio n e f f i c i e n c y
functions were separate and independent physiological systems in wheat
pl ant s.
Both t o ta l
plant nitrogen and nitrogen tran sl oca tio n e f f i c i e n c y
could be used in select ing wheat genotypes f o r e f f i c i e n t nitrogen
u tilizatio n
(Rao et a l . ,
1977).
Studies of c h a r a c t e r i s t i c s rela te d to nitrogen u t i l i z a t i o n provide
useful
information f o r parent selections and planned crosses.
et a l .
(1978) crossed two spring wheat genotypes with complementary
values of t o t a l
efficiency.
high t o ta l
efficiency.
Edwards
reduced plant nitrogen and nitrogen tran sl oca tio n
The high grain protein progeny had a combination of both
reduced plant nitrogen and high nitrogen t ra nsl oca tio n
Bhatia (1975) concluded th at grain protein y i e l d per unit
ground area provides a good selection c r i t e r i o n fo r improving protein
p r o d u c t i v i t y in spring wheat c u l t i v a r s .
McNeal et a l .
(1982) also found
selec tion f o r high grain protein y i e l d increased grain y i e l d and protein
p r o d u c t i v i t y in the progeny of spring wheat crosses.
Chlorophyll Content
The negative association between grain y ie ld and percent grain
protein is due in part to the competition between carbohydrate and
protein accumulation fo r assimilates and energy in plants (Bhatia and
Robson, 1976).
Penning de Vries et a l . (197(0 concluded I g of glucose
produced by photosynthesis in plants can be used to produce O.83 g of
carbohydrate or 0.40 g of protein (assuming n i t r a t e to be the N source).
5
Increasing both grain y i e l d and grain protein content could be achieved
by increasing photosynthetic output, by increasing the ra t e of photosyn­
t h e s i s , and by extending the period of photosynthetic a c t i v i t y (Bhatia
and Robson, 1976).
Cox et a l .
(1985b) found .10 to 22% of the total
plant nitrogen accumulated a f t e r anthesis.
They concluded c u l t i v a r s
with longer green tissue duration a s s im i la t e more nitrogen than c u l t i ­
vars with short green tissu e duration.
Mikesell et a l .
(1971) found
s i m i l a r N content in f l a g leaves o f high and low protein wheat lines at
anthesis.
They reported removing the f l a g leaves at anthesis had l i t t l e
e f f e c t on grain N content of low protein wheat l i n e s , but g r e a t l y
decreased grain N content of high protein wheat l in e s.
They concluded
the v i a b i l i t y and longevity of f l a g leaves are important f o r high
protein l in e s .
anthesis.
High protein lines continued as s im ila ti on of N a f t e r
Neales et a l . (1963) examined the e f f e c t of l e a f removal at
anthesis on grain N content.
They found l e a f removal at anthesis
reduced the N uptake and the grain N content at ma tu rit y.
al.
Spiertz et
( 1971) concluded from 61 to 8 l% of the v a r i a t io n of grain y ie ld
could be s t a t i s t i c a l l y predicted by green area duration of f l a g l e a f and
peduncle.
Rahman (1983) found a p o s it iv e c o r r e l a t io n between net photo­
synthetic ra t e and chlorophyll content per u n i t l ea f area in couchgrass.
6
MATERIALS AND METHODS
Cult ivars
S i x t y - s i x c u l t i v a r s selected from four In te rn ati ona l Winter Wheat
Nurseries (1980, 1981, 1983, 1984) were used in t h is study (Table I ) .
Table I .
Name
Brule
Grana
Bezos. I
AWl2399
Arina
NE7060
Od I ssa-4
A t l a s -66
Lavr in-24
Lethb.32
Martonv.5
Purdue
Blueboy
Houser
Jana
Alcedo
Au ra
MV-6
Martonv.
NS2699
Redwin
Lancota
Names and o r i g in s of c u l t i v a r s used.
Name
Origin
USA,NE
Poland
USSR
E . Germany
Swit z e r .
USA,NE
USSR
USA,IN
Romania
Canada
Hungary
USA,IN
USA,NC
USA,NY
Poland
E . Germany
Finland
Hungary
Hungary
Yugoslav ia
USA,MT
USA,NE
Sutjeska
Lavr in-32
Orov.
F29-75
MV-7
Horosh.
CA8055
NS-15-89A
Katya a - I
Feng-Kang
Saiente
F29-76
GK-Prot.
CIemen t
Doina
WWP.4394
Bastion
TX71A562-6
WW330
Chokwang
Centurk
MT 7811
Origin
Yugoslavia
Roman I a
Yugoslavia
Romania
Hungary
Japan
China
Yugoslavia
Bulgaria
Ch i na
Italy
Romania
Hungary
Netherlands
Romani a
Austria
Netherlands
USA,TX
Australi a
Korea
USA,NE
USA,MT
Name
Vratza
Daws
MV-22-27
Val a
Stephens
Ogosta
WWP.4258
Vega
NS2630-1
Sudova S.
Loudog.
Bounty
Pai Yu P.
NS2699
NE79Y90576
NSR-I
Adams
Inernio
Trakia
Marisml.
PL V
Norwin
Origin
Bulgaria
USA,WA
Hungary
Czechs I .
USA,OR
Bulgaria
Austria
Bulgaria
Yugoslavia
Bulgaria
Bulgaria
England
Ch i na
Yugoslavia
USA,NE
Yugoslavia
Austria
Italy
Bulgaria
England
USA,KA
USA,MT
The c u l t i v a r s were selected to provide a wide range of grain protein
content, grain y i e l d , and genetic backgrounds.
from 20 countries.
The c u l t i v a r s ori gin at ed
They were grouped by height into three groups to
7
reduce i n t e r p l o t competitibn with 22 c u l t i v a r s in each group:
75
- 85cm- - Short
85
~ 95cm- - Medium
>95 cm — T al l
Experimental Design
The t hree height groups were planted separately in a completely
randomized block f i e l d experiment with s i x r e p l ic a te s per harvest in
1984 and 1985 near Bozeman.
nitrogen l e v e l .
spacing.
The groups were randomized wit hi n each
The c u l t i v a r s were planted in h i l l
plots with 0.3 m
T h i r t y seeds of each c u l t i v a r were planted in each h i l l
Seeds were dropped in 3"4 cm deep holes dug with hoes.
plot.
Each r e p l i c a t e
was surrounded by a short winter wheat c u l t i v a r to avoid shading.
Soi I Nitrogen
The soil contained HO and 22 kg/ha N03-N in one meter depth in
1984 and 1985 harvest y e a r s , re sp e c t iv e ly .
three s oi l nitrogen lev el s in 1984 and 1985*
no nitrogen added in both years (NO).
The experiments included
The f i r s t experiment had
The second experiment had 85 and
HO kg/ha nitrogen added as ammonium n i t r a t e (NH^NO^)
and 1985, resp e ct iv e ly ( N i ) .
in 1984
The t h i r d experiment had 170 and 220 kg/ha
NHljNO3 added in 1984 and 1985, re sp e ct iv e ly (N2) .
applied in mid-May.
( 34 - 0- 0 )
F e r t i l i z e r was
These soi l nitrogen levels in two years tota led s i x
soil nitrogen environments.
8
Chlorophyll Estimations
The need to estimate chlorophyll concentrations in f l a g leaves of
f i e l d grown wheat c u l t i v a r s required a nondestructive method of analy­
sis.
Rahman ( 1983) suggested visual color ratings provide an estimate
of chlorophyll content per un i t area.
He found a p o s it iv e c o r r e l a t io n
between visual color ratings and chlorophyll content per un i t area.
A chlorophyll meter s i m i l a r to one used by Wallihan (1973) to
estimate chlorophyll concentrations in leaves o f c i t r u s trees was used
in t h is study.
C a li b r a t io n of the chlorophyll meter included samples of
leaves taken from 17 f i e l d grown wheat c u l t i v a r s .
The leaves were
selected to provide a wide range o f chlorophyll content on the basis of
t h e i r color (dark green to y e l l o w ) .
Light absorbance readings were
taken by chlorophyll meter and chlorophyll concentrations were estimated
by e xt r ac t io n
(Arnon, 1949).
P l o t t i n g chlorophyll concentration (mg/g fresh weight) against
absorbance readings showed 91% of the t o t a l v a r i a t i o n
in chlorophyll
concentration could be s t a t i s t i c a l l y predicted by absorbance readings
taken by chlorophyll meter.
The chlorophyll meter provided a good
estimate of chlorophyll concentration in wheats.
Measurements
Heading dates and plant height were recorded f o r each h i l l
The heading date was the number of days from January I u n t i l
heads in a plot were f u l l y out of the boot.
plot.
50% of the
Plant height was measured
9
in centimeters from the s oi l surface to t i p of the m a jo r it y of spikes
w it h in a p l o t , excluding awns.
Flag l e a f chlorophyll concentration of the 66 c u l t i v a r s was e s t i ­
mated using the chlorophyll meter during grain f i l l i n g period; four
times in 1984 and f i v e times in 1985.
Chlorophyll duration was the
number of days from anthesis u n t i l 75% of f la g leaves in a pl o t turned
yellow.
The following data were recorded f o r each experimental u n i t :
1.
Grain Protein Percentage (GPP) — amount of protein
expressed as percent of the dry weight of the gr ai n.
2.
Grain Yield (GY) — grain weight ( g / p l o t ) .
3.
Biological Yi el d (BY) — t o t a l dry weight ( g / p l o t ) of
aerial
biomass including gr ai n.
4.
Harvest Index (Hi) - - GY/BY.
5.
Total Plant Protein (TPP) — amount of protein
in the a e r i a l
6.
(g/plot)
biomass at mat ur ity including g r a i n .
Grain Protein Yield (GPY) - - amount of protein (g/plot.)
in the grain (GY x GPP).
7.
Nitrogen Harvest Index ( N H l ) " - - GPY/TPP.
Percent protein in the grain (Williams,
1979) and the straw were e s t i ­
mated using the Near Infrared Reflectance analyzer (NIR)
( Noaman et a l . ,
1984).
S t a t i s t i c a l Analyses
Analyses of variance were computed f o r data from each year and
combined over years and soi l nitrogen l e v e l s .
The pooled mean square
10
e r r o r was used to t e s t the c u l t i v a r mean squares and the c u l t i v a r x
environment in te ra c ti o n
fixed.
(McIntosh, 1983)•
Al I factors were considered
Phenotypic c o r r e l a t io n s among t r a i t s were computed f or each year
using entry means over soi l nitrogen l e v e l s .
The r e la tio ns hi ps among
t r a i t s were analyzed using a m u l t i p le regression procedure (Meter and
Vasserman, 131b) .
The rela tio ns hi ps aalong t r a i t s were examined using only c u l t i v a r s
consistent fo r high and low percent grain pr ot ei n.
grain protein i f
A c u l t i v a r had high
i t ranked in the top 10 (of 66) in at le ast four of six
soil N environments.
A c u l t i v a r was considered low grain protein i f i t
ranked in the lowest 10 (of 66) in at least four of s i x soi l N environ­
ments.
The c u l t i v a r s were divided by percent grain protein content into
three groups:
High grain protein c u l t i v a r s
- - >15%
grain
protein
Medium grain protein c u l t i v a r s
— 13-15% grain
protein
Low grain protein c u l t i v a r s
— <13%
protein
grain
RESULTS AND DISCUSSION
Growing conditions d i f f e r e d between the two years of t h is study.
Environmental conditions were ideal f o r wheat growth in 1984.
The soil
nitrogen was high (H O kg/ha) and water stress occurred only l a te in the
grain f i l l i n g
pe r io d .
In the 1985 experiment the soil nitrogen was low
(22 kg/ha) with the drought beginning at heading.
Although i r r i g a t i o n
was applied at heading in 1985» grain y i e l d was about 50% higher in 1984
(Tables 11-16, Appendix).
C u l t i v a r V a ri a t io n in Grain
Protein and Other T r a i t s
The analyses of variance showed highly s i g n i f i c a n t d iff ere nc es fo r
all
t r a i t s among c u l t i v a r s w ithin each year and combined across years.
S i g n i f i c a n t c u l t i v a r x environment i n te ra ct io ns were a t t r i b u t e d l ar ge ly
to years rather than soil N le v e l s .
C u l t i v a r mean squares were much
l ar ge r than c u l t i v a r x environment in te ra ct io ns
(Tables 7 - 10 , Appendix).
In 1984, the check treatment (NO) soil nitrogen content was high (HO
kg/ha).
Nitrogen added to the soi l
(NI and N2) caused no s i g n i f i c a n t
changes in grain y i e l d , biolo gic al y i e l d , and to ta l pl ant p r o t e i n .
Grain protein percentage s i g n i f i c a n t l y increased with increasing soil
nitrogen.
In 1985, the soil nitrogen content was low (22 kg/ha)
check (NO) treatments.
in the
S i g n i f i c a n t d iff ere nc es fo r a l l measured t r a i t s
were obtained among nitrogen l e v e l s .
12
I
PROTEIN X
Figure I .
Frequency d i s t r i b u t i o n of percent grain protein f or 66
c u l t i v a r s at 3 soil N levels (1984).
U
15
PROTEIN %
Figure 2.
Frequency d i s t r i b u t i o n of percent grain protein fo r 66
c u l t i v a r s at 3 soil N levels (1985).
13
This suggested genetic v a r i a b i l i t y
in the e l i t e
in te rn ati ona l
w in te r wheats fo r grain y i e l d , percent grain pr o t e in ,.a n d other t r a i t s
Percent grain protein of the 66 c u l t Ivars was symetrica11y
evaluated.
d i s t r i b u t e d around the means in 1984 f o r a l l
(Figure I ) .
three soi l nitrogen regimes
The low protein c u l t i v a r s increased t h e i r protein content
with increased soi l N, while few changes in grain protein content
occurred in the high grain protein c u l t i v a r s .
Although the analysis of
variance revealed s i g n i f i c a n t d iff er en c es among soil N l e v e l s , the range
of grain protein content was the same at three soil N lev el s (from 11 to
1 8 %).
Percent grain protein o f the 66 c u l t i v a r s in 1985 was symetri c a l I y
d i s t r i b u t e d around individual means f o r each of the three soil N levels
(Figure 2 ) .
pr o t e i n .
Increased soi l N s i g n i f i c a n t l y increased mean percent grain
The range and the v a r i a t i o n f o r percent grain protein were
s i m i l a r at the three soi l N regimes.
The s i g n i f i c a n t di ffe re nc es f o r grain protein content in 1984 and
1985 among c u l t i v a r s and the wide range of grain protein at a l l
levels
soil N
(Figures I and 2) suggested d i f f e r i n g genetic ca p a c i ti e s for
accumulation of protein in the g r ai n .
This is supported by the high
h e r i tab?I i t y of percent grain protein in wheat (Stuber et a l . ,
L o f f l e r and Busch, 1982; Cox et a l . ,
1985a).
1962;
Frequency d i s t r i b u t i o n s of
grain protein content in the. 66 c u l t i v a r s approaching normality at a l l
soil N levels could be an evidence f o r q u a n t i t a t i v e gene control of per­
cent grain protein in these c u l t i v a r s .
14
Relationships Among T r a i t s
Dry■Matter
P o si ti ve c o r r e l a t io n s were found between bio log ical y i e l d and grain
yield
in both years (Table 2 ) .
protein were not c o r r e l a t e d .
Biological y i e l d and percent grain
Thus select ion fo r high bio lo gi c al y ie ld
could improve grain y i e l d without reducing grain pr ot ei n.
Significant
increases in grain y i e l d without reduction in percent grain protein were
achieved by select ion f o r biolo gic al y i e l d
Table 2.
( L o f f l e r et a l . ,
1982) .
Co rrelation c o e f f i c i e n t s ( r ) among t r a i t means o f 66 c u l t i v a r s
over 3 soil N levels in two years.
Biological
Yield
Grain
Yield
Harvest
Index
% Grain
Protein
Total
Plant
Protein
Grain Yield
0.96 sVsV +
0.97 A* (3
Harvest Index
0.14 ns
0.17 ns
0.42 * a
- 0.06 ns
- 0. 0 4 ns
-0 .0 3 ns
-0 .2 7 *
- 0.09 ns
- 0.76 AA
- 0.20 ns
Total Plant
Protein
0.97 * *
0.97 AA
0.91 * *
0.96 AA
0.07 ns
- 0 . 0 7 ns
0.13 ns
0.13 ns
Grain Protein
Yield
0.97 AA
0.95 AA
0.93 * *
0.97 * *
0.15 ns
0.01 ns
0.21 ns
0.16 ns
0,99 * *
0.03 ns
-0 .2 4 ns
• 0.19 ns
- 0.11 ns
0.60 AA
0.52 * *
0.22 ns
- 0.10 ns
-0 .1 7 ns
% Grain
Protein
N Harvest
I ndex
*, **
ns
+
@
0.25 *
= S i g n i f i c a n t at P<0.05 and <0.01, respe ctive ly
= Not s i g n i f i c a n t
=1984
= 1985
0.99 * *
15
The c o r r e l a t io n between grain y i e l d and grain protein percentage
was negative and s i g n i f i c a n t
in 1984, and not s i g n i f i c a n t in 1985
(Table 2 ) .
Consistent high or low grain protein across environments is import­
ant in wheat breeding.
Therefore, the rel ati on sh ip s among t r a i t s were
examined using only c u l t i v a r s consistent for high and low grain p ro te in ,
delet ing the intermediate groups causing an increase in c u l t i v a r x
environment i n te r a c ti o n s .
The grain y i e l d and percent grain protein of
high and low grain protein c u l t i v a r s are plot ted in Figure 3-
PROTON X
Figure 3.
The r e l a t i o n s h ip between grain y i e l d and grain protein (%)
fo r high and low grain protein c u l t i v a r s over 3 soil N levels
(1984).
Despite the negative c o r r e l a t i o n between grain y i e l d and grain
p r o t e i n , c u l t i v a r s with high percent grain protein and reasonably high
16
grain y i e l d were i d e n t i f i e d
(Figure 3 ) .
The simultaneous improvement of
both grain y i e l d and grain protein percentage could be achieved by
se l e c t i o n .
Si milar conclusions were reported by L o f f l e r et a l .
(1985)
and Hal Ioran (1981).
Negative c o r r e l a t io n s between percent grain protein and harvest
index of - 0 . 76* * and - 0.20 were noted in 1984 and 1985, res pe ctively
(Table 2 ) .
High grain protein c u l t i v a r s had low harvest index and vice
versa (Figure 4 ) .
Selection f or high harvest index to improve grain
y i e l d could r e s u l t in reduced grain protein percentage.
Sim ilar high
negative c o r r e l a t io n s between harvest index and grain protein were
e.se-
0.S4-
PR O TD N X
Figure 4.
Grain protein (%) and harvest index for high and low grain
protein c u l t i v a r s over 3 soil N levels (1984).
17
reported by McNea I et a l . (1968), Bhatia (1975), and L o f f l e r et a I .
(1982).
They suggested as HI
increases, the biomass of v eg et a tiv e plant
parts which serve as an N rese rvoir decreases.
T he re fo r e, less nitrogen
is a v a i l a b l e fo r t ra nsl oca tio n to the developing grains.
Plant Protein
Total plant protein accumulated in above-ground parts of the wheat
plants was p o s i t i v e l y cor related with bio lo gi c al y i e l d and grain y i e l d ,
but not cor re la te d with percent grain protein in 1984 and 19&5 (Table
2).
Selection f o r high t o t a l plant protein could increase grain y i e l d
without decreasing grain pr o t e i n .
Total plant protein was strong Iy
associated with both grain y i e l d and grain protein y i e l d accounting fo r
83 to 98% of t h e i r v a r i a t i o n
(r = 0.91 and 0 . 9 9 , re sp e c t iv e ly ; Table 2 ) .
This is in agreement with Neales et a I . ( 1963) and Cox et a l .
(1985b),
Nitrogen harvest index is the genotype's a b i l i t y to p a r t i t i o n
nitrogenous compounds between the grain and the v eg et a tiv e plant parts.
The NHI values from 0.60 to O.83 are w it h in the range reported by Dubois
and Fossati
(1981) f o r winter wheat c u l t i v a r s .
This range was higher
than that reported by Halloran and Lee (1981), and L o f f l e r et a l .
( i g 8 5 ) , fo r spring wheat c u l t i v a r s .
S i g n i f i c a n t p o s it iv e c or r el at io n s
between NHI and harvest index of 0.60 and 0.53 were found in 1984 and
1985, r es pe ct ive ly (Table 2 ) .
This suggests tran slo cat ion of carbohydrate and nitrogenous
compounds are associated.
protein (r = 0.25 P O . 05)
The c o r r e l a t i o n between NHI and percent grain
in 1985 was s i g n i f i c a n t but low, and not
s i g n i f i c a n t in 1984 (Table 2 ) .
Factors other than NHI are important f or
18
fjrain protein percentage.
However, the high grain protein c u l t i v a r s had
higher NHI than the low grain protein c u l t i v a r s
T
r
12
13
t
14
t
1»
I
(Figure 5)•
*
IS
17
Grain yi e l d
t
IS
*
It
PROTEIN X
Figure 5•
N harvest index and grain protein (%) for high and low
grain protein c u l t i v a r s over 3 soil N levels (1985).
was not corr el at ed with NHI
in e i t h e r year.
A greater e f f i c i e n c y for
tran sloc atio n of nitrogen to the grain should resul t in increased grain
protein content at e x i s t in g grain y i e l d le v e l s .
These r es u lt s are in
contrast with those reported in other wheat studies (Cox et a I . ,
L o f f l e r et a l . , 1982).
1985;
They found NHI was p o s i t i v e l y cor re la te d with
grain y i e l d but not corr el at ed with grain p r o t e i n .
They concluded
select ion f or high NHI could increase grain y i e l d without reduction in
grain p r o t e i n .
However, Halloran and Lee (1979) suggested selection fo r
19
high NHI could increase grain protein content without changing grain
yield.
Nitrogen harvest index (%) for high, medium, and low grain protein
c u l t i v a t e at six soil nitrogen levels is shown in Figure 6 .
HIGH GRAIN PROTEIN CULTlVARS
LOW GRAIN PROTEIN CULHVARS
260 +
N O -6 5
N I- 85
N I- 84
N 2 -8 5
N 2 -8 4
SO IL N LEVELS
+ = kg/ha nitrogen ( a v a i l a b l e + added)
Figure 6 .
N harvest index fo r 3 grain protein wheat groups at 6
I evels of soil N.
S im ila r nitrogen harvest index occurred in high, medium, and low
grain protein c u l t i v a r s at low soi l nitrogen le ve ls .
index decreased with increased soil nitro ge n.
c u l t i v a r s were more e f f i c i e n t
Nitrogen harvest
The high grain protein
in t ra nsl oca tin g N to the grain than low
grain prot ein c u l t i v a r s above 110 kg/ha of soi l nitrogen.
Separation of
20
high, medium, and low grain protein c u l t i v a r s on the basis of nitrogen
harvest index could be achieved at any soi l N levels above 110 kg/ha.
High soil nitrogen is required f o r the phenotypic expression of the
genes c o n t r o l l i n g the nitrogen harvest index.
reported by Halloran (1981)
A s i m il a r conclusion was
in a study of nitrogen harvest index of six
spring wheat c u l t i v a r s a t d i f f e r e n t s oi l N le v e l s .
He reported the high
grain protein c u l t i v a r s maintained t h e i r high e f f i c i e n c y in tra ns­
locating N to the grain at high soil N l e v e l s .
He suggested selection
f o r high nitrogen harvest index should be conducted at high soil
nitrogen le v e l s .
Chlorophyll Content
S i g n i f i c a n t d iff ere nc es among c u l t i v a r s f o r f l a g l e a f chlorophyll
concentration (mg/gFW), estimated by a chlorophyll meter, occurred at
anthesis, e a r l y dough, mid-dough, and hard dough in both years and
combined over years (Tables 9 and 10, Appendix).
declined progressively from anthesis to m a t u r it y .
Chlorophyll content
Although the range of
chlorophyll content was very small at anthesis ( 2 . 1 1 to 2.79 mg/g FW),
the range increased at the subsequent stages.
The chlorophyll concen­
t r a t i o n s of 2.11 to 2.79 mg/g FW were w it h in the ranges reported by
Purves and Barmore (1981), Thimann (1985)» and Wallihan ( 1973)•
S i g n i f i c a n t diffe re nc es among c u l t i v a r s f o r chlorophyll duration
were found in both years and combined over years (Table 10, Appendix).
The range was 36 to 49 days in 1984 (Tables 11-13, Appendix) and 30 to
46 days in 1985 (Tables 14-15, Appendix).
S i g n i f i c a n t d iff er en c es among
soil nitrogen levels f o r c h l or op hy ll. c on te nt occurred in both years.
21
Chlorophyll content and chlorophyll duration decreased with added
nitrogen (Tables 17-19» Appendix).
Wide ranges of chlorophyll content
in the f l a g leaves of the 66 c u l t i v a r s were,obtained at e a r l y dough,
mid-dough, and hard dough (Tables 11- 16 , Appendix).
Significant
p o s it iv e c o r r e l a t io n s were found between f l a g l ea f chlorophyll concen­
t r a t i o n at e a r l y dough, mid-dough, and hard dough and bio lo gi c al y i e l d ,
grain y i & l d , t o t a l
1985 (Table 3) •
plant p r o t e i n , and grain protein y i e l d
in 1984 and
S i g n i f i c a n t negative c o r r e l a t io n s occurred between
chlorophyll content and percent grain protein in 1984, but not in 1985
(Table 3)•
Harvest index was p o s i t i v e l y corr el at ed with chlorophyll
content in 1984, but not in 1985.
Table 3•
Co rrelation c o e f f i c i e n t s among t r a i t s of 66 genotype means
over 3 soil N levels in two years.
BY
GY
HI
GPP
TPP
GPY
NHI
C H L O R O PH YLL
C O N C E N T R A T IO N :
E.dough
0.42**+
0.48**@
0.50**
0.51**
0.40**
0 . 15ns
-0.50**
- 0 . 16ns
0.35**
0.49**
0.34**
0.48**
- 0 . 06ns
-0.27*
M.dough
0.45**
0.4?**
O. 51* *
0.49**
0. 32 *
0 . 12ns
-0.49**
- 0-. 15ns
0.37**
0.49**
0.35**
0.46**
- 0 . 13ns
- O . 39* *
H.dough
0.40**
0.39**
0.46**
0.40**
0. 30 *
0 . 05ns
-0.41**
- 0 . 09ns
0. 33 *
0.42**
0.32*
0.38**
- 0 . 06ns
- O . 39* *
- 0 . 18ns
0.29*
- 0 . 22ns
0 . 28*
- 0 . 23ns
0 . 05ns
O. 39* *
0 . 07ns
- 0 . 07ns
0.29*
- 0 . 09ns
0.31*
- 0 . 18ns
0 . 13ns
D U R A T IO N :
BY = Biological Y i e l d ; GY = Grain Y i e l d ; HI = Harvest Index; GPP = Grain
Protein Percentage; TPP = Total Plant P r o t e i n , GPY = Grain Protein
Y i e l d , NHI = Nitrogen Harvest Index
* , * * = S i g n i f i c a n t at P<0.05 and P<0.01, respe ct ive ly
ns = Not s i g n i f i c a n t
+ = 1-984; @ -
1985
22
Grain y i e l d was more p o s i t i v e l y associated with chlorophyll concen­
tration
content.
in the f l a g l e a f during grain f i l l i n g
than with grain protein
. Most of the carbohydrate a v a i l a b l e f o r grain f i l l i n g
is
dependent on size and duration of green tissu e during the grain f i l l i n g
period.
Only 5 to 10% of the carbohydrate formed before anthesis is
a v a i l a b l e fo r r e d i s t r i b u t i o n to the grain (Lupton, 1968).
However,
nitrogen is mostly accumulated in the v eg et a tiv e plant parts p r i o r to
anthesis and then translocated to the g r a i n .
Only 10 to 22% of the
t o t a l nitrogen is assimilated during the grain f i l l i n g
a I . , 1985b).
period (Cox et
The photosynthesis of the f l a g l e a f in wheat makes a major
con tri bu tio n of the carbohydrate to the developing grain (S p i e r tz et a l ,
1971).
The negative association between chlorophyll content and grain
protein is i n d i r e c t evidence of the p o s i t i v e association o f breakdown of
chlorophyll and l e a f pr o t e i n .
Peoples et a l .
(1980) reported ribulose
bisphosphate carboxylase (RuBPCase) was the major l e a f storage protein
as well as a major c a t a l y s t of CO2 f i x a t i o n .
of the t o t a l
soluble l e a f protein in wheat.
associated with degradation of chlorophyll
RuBPCase accounted f o r 60%
RuBPCase degradation is
(Hall et a l , 1978).
Percent
grain protein was not corr el at ed with chlorophyll concentration at three
grain f i l l i n g
stages in 1985 (Table 3 ) .
Drought stress e a r l y in the
growing season which led to e a r l y heading and ea r ly m a tu r ity may have
been a f a c t o r .
Nitrogen harvest index was neg atively cor re la te d with
chlorophyll concentration at mid-dough and hard dough in 1985, but not
in 1984 (Table 3 ) .
This suggests high percent grain protein was not
associated with higher concentration of chlorophyll
at any growth stage.
in the f l a g leaves
Si m ila r conclusions were reported by Cox et a l .
23
(1985a).
They found that c u l t i v a r s with lower percent green tissue
(estimated k~] days a f t e r anthesis)
translocated more N to the grain than
c u l t i v a r s with higher percent green tiss ue .
Chlorophyll duration estimated from anthesis to the complete
yellowing of the fla g l e a f was s i g n i f i c a n t l y corr el at ed with bio log ical
y i e l d and grain y i e l d
in 1985 (r = 0.29 and 0.28 P<0.05, r e s p e c t i v e l y ) ,
but not in 1984 (Table 3 ) .
Grain protein percentage was p o s i t i v e l y
corr el at ed with chlorophyll duration in 1984 (Table 3) , but not in 1.985 •
Although the c o r r e l a t io n c o e f f i c i e n t between chlorophyll duration and
grain prot ein was low, the high grain pr otein c u l t i v a r s had longer
chlorophyll duration than the low grain protein c u l t i v a r s
(Figure 7)•
The chlorophyll duration f o r high and low grain protein c u l t i v a r s at
three soi l N levels f o r two years is shown in Figure 8.
The high grain
protein c u l t i v a r s had longer chlorophyll duration than low grain protein
c u l t i v a r s at any level of soi l nitrogen in both 1984 and 1985.
This
indicates long chlorophyll duration is associated with high grain
protein c u l t i v a r s .
The importance of the longevity of f l a g leaves and high protein
wheat was reported by Neales et a I . ( 1963) and Johnson e t a I . (1968).
They concluded the rete nt ion of leaves is important f o r phenotypic
expression of high protein in wheat.
chlorophyll
Diseases that damage the
in the leaves would adversely a f f e c t the level of prot ein .
Close association of l e a f rust resistance and high prot ein in wheat was
reported by Haunold e t a l .
(1962).
24
Figure 7•
Chlorophyll duration and grain protein (%) fo r high and low
protein c u l t i v a r s over 3 soil N levels (1984).
HIGH GRAIN PROTEIN CULTIVARS
LOW GRAIN PROTEIN CULTIVARS
NI
SOIL N LEVELS
Figure 8.
Chlorophyll duration for high and low grain protein groups at
3 soil N levels in two years.
25
Grain Yield and Grain Protein
as Functions of Several T r a i t s
Percent grain protein and grain y i e l d as an expression of several
t r a i t s was examined using m u l t i p l e regression.
The pred ic tio n of GPP
including GY, NHI, and TPP as independent var ia bl es accounted for 93%
and 90% of the t o t a l v a r i a t i o n o f GPP among c u l t i v a r s in 1984 and 1985,
r es pe ct ive ly (Tables 4 and 5 ) .
Both NHI and TPP had p o s i t i v e regression
c o e f f i c i e n t s , whil e GY produced a negative regression c o e f f i c i e n t .
given grain y i e l d l e v e l ,
Table 4.
At a
increasing both NHI and TPP could increase GPP.
M u l t i p l e regression analyses expressing percent grain protein
and NHI as functions of several t r a i t s (1984).
Independent
Variables
Grain
Protein (%)
GY,TPP,NHI
NHI
GY,TPP,GPP
NHI
GPY,SPY
. Regression Co e ff ic ie n ts
BI
B2
B3+
R2
-0.28
1.56
**
15-40
0.83
0.06
0.03
ft*
0.60
—
0.93
O
Dependent
Va riable
Aft
I
m
cU :
O
0.09
ft*
A*
GY = Grain Y i e l d ; TPP = Total Plant Pro tein ; NHI = Nitrogen Harvest
Index; GPY = Grain Protein Y i e l d ; SPY = Straw Protein Yi el d
+ =Bl,
B2, B3 = M u l t i p l e regression c o e f f i c i e n t s
* * = S i g n i f i c a n t at P<0.01
The pr edict ion of NHI including GY, TPP, and GPP (Tables 4 and 5)
as independent var ia bl es accounted f o r 60% and 73% o f the t o t a l v a r i a ­
tion among c u l t i v a r s in 1984 and 1985, re sp e ct iv e ly .
26
Table 5.
M u l t i p l e regression analyses expressing percent grain protein
and NHI as functions of several t r a i t s (1985).
Dependent
Va riable
Independent
Variables
Grain
Protein (%)
GY,TPP1NHI
NHI
GY1TPP1GPP
Regression Co e ff ic ie n ts
BI
B2
BB+
R2
-0 .8 1
4.40
18.20
0.90
AA
AA
0.03
AA
GPY1SPY
NHI
0.06
AA
0.04
- 0 . 18
0.73
AA
AA
—
.- 0 . 2 1
0.91
AA
GY = Grain Y i e l d ; TPP = Total Plant Protein; NHI = Nitrogen Harvest
Index; GPY = Grain Protein Y i e l d ; SPY = Straw Protein Yiel d
+ = BI, B2, BB = M u l t i p l e regression c o e f f i c i e n t s
* * = S i g n i f i c a n t at P<0.01
Grain y i e l d gnd percent grain protein had p o s it iv e regression c o e f f i ­
c i e n t s , while t o t a l
plant protein had a negative regression c o e f f i c i e n t .
The pr edi ction of NHl using only grain protein y i e l d
straw protein y i e l d
(GPY) and
(SPY) accounted f o r 93% and 91% of the t o ta l v a r i a ­
tion in 1984 and 1985, resp e ct iv e ly (Tables 4 and 5 ) .
Grain protein
y i e l d had a p o s it iv e regression c o e f f i c i e n t , while amount o f protein
remaining in the straw had a negative regression c o e f f i c i e n t .
This
suggests that high e f f i c i e n c y of tra ns lo ca tin g N from the straw to the
grain is associated with high grain y i e l d and high percent grain
protein.
M u l t i p l e regression equations expressing percent grain protein and
grain y i e l d as functions o f several t r a i t s are shown in Table 6.
The
prediction, of percent grain protein included t o ta l plant p r o t e i n , NHI,
27
chlorophyll du rat io n, and grain y i e l d .
This combination of independent
v a r i a b l e s accounted f o r 94% and 90% of the t o t a l v a r i a t i o n
grain protein in 1984 and 1985, re sp e c t iv e ly .
in percent
The grain y i e l d predic­
tion equation including t o t a l plant p r o t e i n , NHI, and chlorophyll dura­
tion accounted f o r 88% and 94% of the t o t a l v a r i a t i o n
1984 and 1985, re sp e c t iv e ly .
in grain y i e l d in
Total plant pr o t e i n , NHI, and chlorophyll
duration were the most important va r i a b l e s in a l l
pred ic tio n equations.
None of the other t r a i t s measured met the required s i g n i fi c a n c e levels
f o r entry into the equations.
Table 6.
M u l t i p l e regression analyses expressing percent grain protein
and grain y i e l d as functions o f several t r a i t s f o r 66
c u I t ivars.
Independent
Variables+
Dependent
Variable
2
Rz
1984
Grain Protein (%)
Grain Yield
TPP, NHI, ChL.D., GY
TPP, NHI , ChL.D.
0.94
0.88
1985
Grain Protein (%)
Grain Yield
TPP, NHI, ChL.D., GY
TPP, NHI-, ChL.D.
0.90
0.94
+TPP = Total Plant Protein; NHI = Nitrogen Harvest Index; ChL.D.
Chlorophyl I Duration; GY = Grain Yield
Selection fo r high NHI, high t o t a l
plant p r ot ei n, and long
chlorophylI duration could increase both grain y i e l d and grain protein
percentage.
Johnson et a l .
(1967) reported nitrogen uptake and n Itrogen
p a r t i t i o n i n g function as separate and independent physiological systems
in the wheat p l a n t .
Both NHI and t o t a l
plant protein independently
contributed to percent grain protein in the current study.
Therefore,
28
inclusion of both characters in wheat genotypes through breeding should
r e s u l t in higher grain y i e l d and higher grain protein content.
Rao et
a I . (1977) suggested that selection f o r e f f i c i e n t nitrogen u t i l i z a t i o n
must include two or more factors simultaneously, such as high N uptake
and high e f f i c i e n c y of tran sl oca tin g ve g et a tiv e nitrogen to the grain.
29
SUMMARY
The low negative c o r r e l a t i o n between grain y i e l d and percent grain
protein in t h i s study indicates simultaneous increases in grain y ie ld
and protein content could be achieved by se l e c t i o n .
analyses suggested th at increases in t o t a l
plant protein may increase
grain y i e l d without reducing percent grain p ro te in .
content was associated with high NHI.
Simple c o r r e l a t io n
High grain protein
Selection f o r high NHI at high
soil nitrogen could increase percent grain protein without reducing
grain y i e l d .
Although s i g n i f i c a n t c o r r e l a t io n s were found between grain y i e l d ,
biol og ic al y i e l d , t o t a l plant protein and chlorophyll concentration in
the f l a g leaves at three grain f i l l i n g
fo r p r e d i c t i v e purposes.
grain prot ein content.
stages, the values were too low
Chlorophyll duration was associated with high
Separation of high and low grain protein
c u l t i v a r s was possible by chlorophyll duration.
The m u l t i p l e regression analyses suggested t o t a l pla nt p ro te in ,
NHI, and chlorophyll duration contributed to both high grain y i e l d and
high percent grain p ro te in .
Combining high NHI, high t o t a l
plant
p r o t e i n , and long chlorophyll duration in h yb ri d iz at io n could provide
wheat c u l t i v a r s with high y i e l d and high protein content.
30
LITERATURE CITED
31
LITERATURE CITED
Arnon, D . 1 . 1949.
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Copper enzymes in isolated chlor opl ast s.
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Austin, R.B., M.A. Ford, J.A. Edrich, and R D. Blackwel I .
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1976.
B io e n e r g e t ic c o n s id e r a tio n m
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Science 194:1410-1421
B h a tia ,
Cox, M.C., C.O. Qua I s e t , and D.W. Rains.
1985a.
Genetic v a r i a t i o n fo r
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I . Dry matter
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Crop S c i . 25:430-435«
Cox, M .C . , C.O. Qua I s e t , and D.W. Rains.
1985b. Genetic v a r i a t i o n for
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I I . Nitrogen
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Crop S c i .
25:435-440.
Desai, R.M., and C.R. Bhatia.
1978.
Nitrogen uptake and nitrogen
harvest index in durum wheat c u l t i v a r s varying in t h e i r gram
protein concentration.
Euphytica 27:561-566.
Dubois, J . B . , and A. F o s s a t i . 1981.
Influence of nitrogen uptake and
nitrogen p a r t i t i o n i n g e f f i c i e n c y on grain y i e l d and grain protein
concentration of twelve win te r wheat genotypes { T r i t i c u m a e s t i v u m
L.).
Z . P f lanzenzucht. 86:41-49.
Edwards, I . B . , J.A. Mey, and M. van der Mey.
1978.
Use of a physio­
logic model f o r g e n e t i c a l l y improving grain protein in wheat.
Cereal Foods World 23=596-600.
H a l l , N . P . , A.J. Keys, and M.J. M e r re tt .
1978.
Ribulose 1,5 diphos­
phate carboxylase protein during f l a g l e a f senescence.
J. Exp.
Bot. 29:31-37H a l l o r a n , G.M.
198I . C u l t i v a r d if f er en c es in nitrogen t ra nsl oca tio n in
wheat. Aust. J. A g r i c . Res. 32:535-544.
Ha ll o r a n , G.M., and J.W. Lee.
1979.
Plant nitrogen d i s t r i b u t i o n
wheat c u l t i v a r s .
Aust. J. Agric. Res. 30:779-789«
in
32
Haunold, A . , V. A. Johnson, and J.W. Schmidt.
1962. V a ri a t io n in
protein content of the grain of four v a r i e t i e s of T f i t i c u m a e s t i v u
L. Agron. J. 5^:121-125.
m
Johnson, V . A . , P.J. Mattern, and J.W. Schmidt.
1967Nitrogen r e l a ­
tions during spring growth in v a r i e t i e s of T r i t i c u m a e s t i v u m L.
d i f f e r i n g in grain protein content.
Crop S c i . 7:664-66?.
Johnson, V . A . , J.W. Schmidt, and P.J. Mattern.
1968.
f o r b e t t e r protein impact.
Econ. Bot. 22:16-25.
Cereal breeding
L o f f l e r , C.M., and R.H. Busch.
1982.
Selection fo r grain p ro te in ,
grain y i e l d , and nitrogen p a r t i t i o n i n g e f f i c i e n c y in hard red
spring wheat.
Crop S c i . 22:591~595«
L o f f I e r , C.M., T.L. Rauch, and R.H. Busch.
1985. Grain and plant
protein rela tio ns hi ps in hard red spring wheat.
Crop S c i .
25:521-524.
Lupton, F.G.H.
1969.
Estimation of y i e l d in wheat from measurement of
photosynthesis and tran sl oca tio n in the f i e l d .
Ann. Appl. Biol.
64:363-374.
McIntosh, M.S.
1983.
75:153-155.
Analysis of combined experiments.
Agron. J.
McNeal, F . H . , G.O. Boatwright, M.A. Berg, and C.A. Watson.
1968.
Nitrogen in plant parts of seven spring wheat v a r i e t i e s at
successive stages of development.
Crop S c i . 8:535-537.
McNeal, F . H . , C.F. McGuire, and D.L. Klindworth.
1982. Agronomic and
q u a l i t y c h a r a c t e r i s t i c s of spring wheat lines selected f o r protein
content and protein y i e l d .
Euphytica 31=377-381.
Mikesel I , M.E., and G.M. Paulson.
1971.
Nitrogen t ra nsl oca tio n and the
role of individual leaves in protein accumulation in wheat grain.
Crop S c i . 11:919-922.
Neales, T . F . , M.J. Anderson, and I . F . Wordlaw.
1963. The ro le of the
leaves in the accumulation of nitrogen by wheat during ear develop­
ment. Aust. J. Agric. Res. 14:725-736.
N e t e r , J . , and W. Wasserman.
1974. Applied l i n e a r s t a t i s t i c a l models.
Regression, analysis of variance, and experimental designs.
Richard D. Ir w i n , Inc.
Homewood, I l l i n o i s .
Noaman, M.M., G.A. T ay lo r , and C.F. McGuire.
1984. The use of NIR f o r
estimation of N/ protein at various growth stages in wheat.
American Soc. Agronomy Ab s tr . November 25-30, Las Vegas, NV.
33
Penning de V r i e s , F . W . T . , A.H.M. Brunsting, and H.M. van Laar.
1974.
Products requirements and e f f i c i e n c y of biosynthesis: A q u a n ti t a ­
t i v e approach.
J . Theor. B i o l . 45:339~377Peoples, M.B., V.C. B e il h a r z , S.P. Waters, R.J. Simpson,and M.J. _
B a ll in g .
I960.
Nitrogen r e d i s t r i b u t i o n during grain growth in
wheat { T r i t i c u m a e s t i v u m L . ) .
I I . Chloroplast senescence and the
degradation of ribulose-1,5~bisphosphate carboxylase.
Planta
149:241-251 .
Purvis, A . C., and C.R. Barmore.
1981.
Involvement of ethylene in
chlorophyll degradation in peel of c i t r u s f r u i t s .
Plant Physiol.
68:854-856.
Rahman, M.S.
1983. Relationship between visual colour r at in g and
chlorophyll content, photosynthetic r a t e , and some growth
c h a r a c t e r i s t i c s in couchgrass ( c y n o d o n s p p . L . ) .
J. Agric. S c i .
100:221-225.
Rao, K . P . , D.W. Rains, C.O. Qualset, and R.C. Huffaker.
1977- Nitrogen
n u t r i t i o n and grain protein in two spring wheat genotypes d i f f e r i n g
in n i t r a t e reductase a c t i v i t y .
Crop S c i . 17:283-286.
S p i e r t z , J . H . J . , B.A. ten Hag, and L . J . P . Kupers.
1971.
Relation
between green area duration and grain y i e l d in some v a r i e t i e s of
spring wheat.
Neth. J . Agric. S c i . 19:211-222.
Stuber, C.W., V.A. Johnson, and J.W. Schmidt.
1962.
Grain protein
content and i t s re la tio ns hi ps to other plant and seed characters in
the parents and progeny of cross of T r i t i c u m a e s t i v u m L. Crop S c i .
2 : 506- 508.
Terman, G . L . , R.E. Ramig, A . F. D r e i e r , and R.A. Olson.
1969- Y i e l d protein rela tio ns hi ps in wheat grain as af fe cte d by nitrogen and
water.
Agron. J. 61:755-759.
Thimann, K.V.
1985. The senescence of detached leaves o f tropaeolum.
Plant Physiol. 79:1107-1110.
Wallihan, E.F.
1973.
Portable r e f le c ta n ce meter f o r estimating
chlorophyll concentrations in leaves.
Agron. J. 65:659"o62.
W ill ia m s, P.C.
1979.
Screening wheat f o r protein and hardness by near
IR r ef le c ta nc e spectroscopy.
Cereal Chem. 56:169-172.
APPENDIX
35
Table 7.
Mean squares of grain y i e l d and percent grain protein of three
height groups of c u l t i v a r s combined over years and soil n i t r o ­
gen le v e l s .
Percent Grain Protein
Grain Yield
d .f.
Source
I
Year(Y)
Nitrogen (N)
2
YxN
2
\
Short
Tal I
Medi urn
178200
131.6
48.61
140.5
AA
AA
AA
AA
63.4
68.8
115.8
AA
AA
AA .
37.4
21.6
41.7 .
AA
AA
AA
0.53
0.52
Tal I
Medi urn
Short
193300
**
187300
**
284.9
ns
294.3
ns
56.2
ns
134.3
ns
41.0
ns
130.9
ns
0.564
BLK/N/Y
30
118.7
142.3
111.0
Cult ivqr (C )
'
21
1124
**
2694
2202
54.0
36.8
29.6
A*
AA
AA
AA
AA
YxC
21
42
NxC
Y x N x C
Pooled Error
ns
42
630
390.6
**
1294
780.6
6.8
7-4
6.4
AA
AA
AA
AA
AA
72.4
ns
99.9
ns
129.8
ns
0.89
I .76
1.9
AA
AA
AA
106.3
ns
89.1
ns
133.0
ns
0.99
1.3
1.21
AA
AA
AA
96.06
88.21
88.26
0.369
0.273
0.238
S i g n i f i c a n t a t P<0.01
Not s i g n i f i c a n t
36
Table 8.
Mean squares of bio lo gic al y i e l d and t o ta l plant protein of
three height groups of c u l t i v a r s combined over years and soil
nitrogen le ve ls .,
Biological Yield
d .f.
Source
T al l
Percent Grain Protein
Medi urn
Short
Tal l
Medi urn
Short
I
1373000
ftft
114000
ft*
998800
ft*
5936
ft*
5333
**
4505
ft*
Nitrogen (N)
2
233.4
ns
4343
ftft
488.1
ns
2.46
ns
30.3
ftft
11.9
ft
YxN
2
4203
ftft
1335
ns
846.9
ns
17.9
ft
2.7
ns
6.7
ns
Year(Y)
■
BLK/N/Y
30
696.4
657-5
859.8
5.0
3.9
3.5
Cult i v a r ( C )
21
4809
ft*
12910
ft*.
110600
ftft
36.8
ft*
68.1
Aft
66.4
ft*
YxC
21
1879
ft*
5542
ftft
3851
**
17.4
ft*
26.8
ft*
25.2
Aft
NxC
42
392.1
ns
568.4
ns
544.6
ns
2.4
ns
3.1
ns
3.9
ftft
Y x N x C
42
441.8
ns
505.3
ns
552.1
ns
3.2
ns
3. 0
ns
4.3
ft*
630
476.7
428.0
416.8
2.65
2.47
2.38
Pooled Error
Af
ns
ft f t
=
=
S i g n i f i c a n t at P<0.05 and P<0.01, res pe ct iv e ly .
Not s i g n i f i c a n t
37
Table 9.
Mean squares of chlorophyll concentration (mg/g FW) at
anthesis and e a r l y dough of three height groups of c u l t i v a r s
combined over years and soil nitrogen le ve ls .
Chlorophyll Concentration
Anthesis
Source
Year (Y)
Nitrogen (N)
d .f.
I
2
2
YxN
Tal I
Medi urn
____
Short.
________ E. Dough
Medi urn
Short
1.42
ns
0.052
ns
0.103
ns
Tal I
11.93
17.79
18.42
A*
AA
AA
0.582
**
0.18
0.475
17-9
25.4
37-7
AA
AA
AA
AA
AA
0.635
0.439
0.301
6.84
9.29
6.16
AA
AA
AA
AA
AA
AA
BLK/N/Y
30
0.0534
0.029
0.031
0.27
0.497
0.337
C u l t i v a r s (C)
21
0.2273
0.274
0.347
2.25
2.85
3-75
AA
AA
AA
AA
AA
AA
0.019
ns
0.068
0.90
0.792
0.727
1.13
AA
AA
AA
AA
AA
0.0196
ns
0.0185
ns
0.0265
ns .
0.318
0.326
0.364
AA
AA
AA
0.0221
ns
0.0313
ns .
0.0236
ns
0.137
0.238
0.274
AA
AA
AA
0.0224
0.0239
0.0233
0.0626
0.10
0.07
21
YxC
NxC
42
Y x N x C
42
Pooled Error
**
ns
630
S i g n i f i c a n t at P<0.01
Not s i g n i f i c a n t
38
Table 10.
Mean squares of chlorophyll concentration at hard dough and
chlorophyll duration of three height groups of c u l t i v a r s
combined over years and soil nitrogen le v e l s .
Chlorophyll
d .f.
Source
I
Year (Y)
Nitrogen (N)
YxN
2
2
Medi urn
Short
2190
AA
4558
AA
5208
AA
1.74
554.7
243-5
548.5
AA
AA
AA
AA
9.34
112.3
241.9
AA
AA
AA
4.58
6.45
4.66
Medium
Short
4.21
3-57
AVe
AA
1. 6 8
*
2.78
4.60
AA
AA
8.88
AA
8.81
AVe
Tall
Chlorophyll Duration
(mg/g FW)
T al l
.
. 113.6
AA
BLK/N/Y
30
0 .2 58
0.229
0.225
Cu lti v ar s( C )
21
4.01
4.86
6.77
145.2
64.2
97.6
AA
AA
AA
AA
AA
AA
0.877
1.13
0.814
40.1
AA
AA
AA
81.2
AA
75.2
AA
0.183
0.273
22.6
AA
37.2
AA
0 .2 18
AA
26.5
AA
AA
AA
0.311
0.265
0.384
24.7
29.7
32.5
AA
AVe
AA
AA
AA
AA
0.0843
0.0739
0.0714
4.61
4.69
4.7
YxC
21
42
NxC
42
Y x N x C
Pooled Error
*.
AVc
630
S i g n i f i c a n t at P<0.05 and P<0.01, re sp e ct iv e ly .
AA
39
Table 11.
Eight t r a i t means of 22 win te r wheat c u l t i v a r s
grown at 3 soi l N levels in 1984.
( t a l l group)
Chlorophyll Concent.
E.
Dough
M.
Dough
H.
Dough
ChL.D.
TPP
NHI
8.1
11.6
9.0
10.4
10.0
5.1
10.4
. 7-3
9. 8
6.9
?.l
7-8
7-3
7. 9
7-1
9. 0
10.8
8. 6
10.3
10.7
0.74
0.76
0.78
0.77
0.78
0.79
0.77
0.77
0.81
0.78
0.75
0.74
0.80
0.78
0.76
0.77
0.76
0.74
0.75
0.77
0.73
0.75
1.92
2.30
1.91
2.24
2.22
1.48
I .94
1.72
2.18
0.74
2.00
I .82
1.92
2.01
2.30
2.15
1.54
2.10
1.95
I .41
2.17
1.95
1.38
1.30
0.77
1.18
1.10
0.32
0.81
0.69
1.10
0.16
0.85
0.65
0.83
0.89
1.34
1.12
0.50
1.00
1.04
0.46
1.26
0.96
0.145
0.561
0.178
0.324
0.299
0.035
0.068
0.067
0.218
0.051
0.110
0.011
0.262
0.257
0.710
0.388
0.305
0.280
0.196
0.255
0.419
0.253
44.5
41.8
41.8
42.2
40.7
42.3
41.6
40.2
40.6
44.4
42.3
43.2
42.3
41.7
41.1
40.0
37.4
41.3
42.8
42.8
44.8
46.2
13-9
9-0
0.77
1.90
0.89
0.245
42.1
19.0
3.2
17^6
5. 0
LSD.05
5.2
0.2
0. 9
0.02
0.12
0.12
0.076
1.0
LSD.Ol
7.3
0.3
1.2
0.30
0.16
0.17
0.107
1.3
GY
GPP
Brule (USA,NE)
Grana (Poland)
Bezost. I (USSR)
AW12399 (E.Germ.)
Arena ( S w i t z e r l . )
NE7060 (USA,NE)
Odi ssa-4 (USSR)
Atl as- 66 (USA,IN)
Lavrin24 (Romania)
Lethb.32 (Canada)
Martonv.5 (Hungary)
Purdue (USA,IN)
Blueboy (USA,NC)
Houser (USA,NY)
Jana (Poland)
Alcedo (E.Germ.)
Aura (Finland)
MV-6 (Hungary)
Mar tonv. (Hungary)
NS2699 (Yugoslav.)
Redwin (USA,MT)
Lancota (USA,NE)
50.2
59.2
55.3
62.4
47-5
49.4
49.9
25.4
60.0
38.2
50.7
37.8
56.6
54.5
47-2
48.2
41.5
49.0
59.0
49.2
49.4
51.4
12.7
12,5
14.0
14.4
14.9
16.7
15.4
15.3
14.1
14.9
14.5
13.5
12.8
11.1
11.7
12.1
13.0
13-7
13.8
13.5
15.2
15.7
Means
49-6
C.V.
C u lti v ar s
9.7
9.9
11.1
7-5
18.7
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
Protein; NHI = Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
4.1
Table 12.
Eight t r a i t means of 22 w in te r wheat c u l t i v a r s (medium
height group) grown at 3 soil N levels in 1984.
Chlorophyll Concent.
GY
GPP
Sutj esk a(Yugos.)
Lavrin^32(Romania)
Orov.(Yugoslavia)
F29~75(Romani a)
MV-7(Romania)
Horosh(Japan)
CA8055(China)
NS-15-89A(Yugosl.)
K a t i a - I (Bulgaria)
Feng-Kang(China)
Saiente(Ita ly )
F29-76(Romania)
GK-Prot. (Hungary)
Clement(NetherId.)
Doina(Romania)
WWP.4394(Austria)
Bastion(Nether I d .)
TX71A562-6(USA,TX)
WW330(Australia)
Chokwang(Korea)
Centurk(USA,NE)
MT 7811(USA,MT)
51.7
45.3
21.7
64.2
49.5
62.0
47.3
39.1
41.1
33.1
38.7
51.8
39.4
55.5
45.7
41.3
23.4
69.1
27.0
13.8
1-3.3
14.2
13.1
13.1
13.4
13.8
14.0
Cu lti va rs
TPP
NHI
E.
Dough
M.
Dough
H.
Dough
ChL.D,
66.0
17.2
13.1
12.9
16.2
11.2
13.0
14.7
12.5
11.9
13.1
15.6
13.8
13.3
9.4
7.5
4.1
10.9
8.3
10.6
. 8.4
7.2
7-3
7.6
6.5
. 8.5
8. 0
8.1
7-5
7-7
3.9
10.2
4.9
6.1
12.0
11.1
0.76
0.80
0.76
0.76
0.78
0.79
0.78
0.75
0.76
0.74
0.78
0.78
0.80
0.77
0.79
0.79
0.75
0.80
0.73
0.74
0.79
0.79
2.07
I .90
1.55
2.15
1.96
2.15
1.82
1.95
1.88
1.15
1.63
2.20
1.53
2.35
1.82
1.89
1.99
1.74
1.93
0.88
1.97
2.28
0.292
1.02
0.69
0.103
0.058
0.34
1.12
0.236
0.75
0.045
1.04
0.275
0.61
0.045
0.83 ■ 0.099
0.134
0.79
0.26
0.058
0.045
0.49
1.00
0.281
0.080
0.32
1.010
1.43
0.75
0.171
0.265
0.79
0.316
1.00
0.149
0.55
0.149
0.77
0.20
0.087
0.380
0.90
1.26
0.477
41.5
43.5
43.6
42.2
42.9
43.3
44.8
43.0
45.3
46.8
42.2
43.1
44.5
42.4
45.7
45.7
44.0
43.9
44.6
46.3
42.2
44.4
Means
45-9
13-7
8. 0
0.77
1.85
0.77
0.216
43.9
C.V.
20.0
2.9
19.1
5.6
LSD.05
5. 0
0.2
0. 8
0.02
0.12
0.11
0.062
1.0
LSD.Ol
7-1
0.3
I .2
0.03
0.17
0.15
0.088
1.3
29.0
66.9
13.6
11.6
8.8
1,6.7
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
P r o t e i n ; NHI = Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
4.0
Table 13.
Eight t r a i t means of 22 w in te r wheat c u l t i v a r s
grown at 3 soil N levels in 1984.
(short group)
Chlorophyll Concent.
Cult i v a r s
TPP
M.
Dough
H.
Dough
0.22
1.16
0.65
0.94
1.02
1.04
0.87
0.20
0.73
I .00
1.10
1.20
0.20
0.56
0.13
0.55
1.20
0.011
0.329
0.156
0.251
0.238
0.293
0.144
0.79
0.79
0.81
1.34
2.25
1.78
2.08
2.14
2.14
1.97
1.31
1.95
2.09
2.24
2.24
1.05
1.57
0.68
1.71
2.06
1.81
I .92
2.46
1.20
2.19
0.79
1.83
NHI
GY
GPP
33.6
59.2
5.2
8.4
8. 8
8.4
7-2
9.2
7-3
6.1
10.2
8.2
13-6
6. 8
7-3
8.9
6.4
5.7
7. 6
3.1
7.4
5.7
8.5
9. 9
0.82
0.80
0.76
0.79
0.82
0.79
0.81
0.80
0.79
0.79
0.75
0.80
0.79
0.78
0.83
0.77
0.82
0.76
E.
Dough
ChL.D.
0.77
1.36
0.36
1.16
0.011
0.278
0.054
0.084
0.369
0.226
0.443
42.7
42.5
42.1
42.4
40.9
44.6
41.9
43-9
42.8
43.6
44.3
40.3
46.3
42.6
43.7
44.8
42.9
44.0
44.0
41.5
45.6
42.2
0.77
0.196
43.2
Vratza (Bulgaria)
Daws (USA1WA)
MV-22-27 (Hungary)
Vala (Czechslov.)
Stephens (USA1OR)
Ogosta (Bulgaria)
WWP.4258 (Austria)
Vega (Bulgaria)
NS2630-1 (Yugosl.)
Sudova S. (Bulgar.)
Loudog. (Bulgaria)
Bounty (England)
Pai Yu P. (China)
NS2699 (Yugoslav.)
NE79Y90576 (USA1NE)
NSR-I (Yugoslavia)
Adams (Austria)
Inernio ( I t a l y )
Trakia (Bulgaria)
Marisml. (England)
PL V (USA1KA)
MT 7877 (USA1MT)
53.1
48.9
55.1
49.4
34.3
58.6
50.2
71.9
43.7
37.7
51.7
40.0
31.0
48.2
18.3
44.9
35.5
38.6
61.1
13-4
11.4
13.2
12.5
12.1
13.2
12.0
14.1
13-7
12.7
14.0
12.5
15.2
13.4
13-2
14.2
13-0
12.9
13-3
12.8
17.3
13.1
Means
41.2
13.3
7.7
C.V.
22.2
3.0
19.5
LSD.05
5.0
0.2
0.8
0.02
0.12
0.11
0.056
1.0
LSD.01
7-1
0.3
1.2
0.03
0.18
0.15
0.079
1.4
50.6
0.80
4.0
12.4
0.71
9-3
0.050
0.120
0.192
0.466
0.363
0.033
0.186
0.030
16.4
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
Protein; NHI = Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
4.1
42
Table 14.
Eight t r a i t means o f 22 w int er wheat c u l t i v a r s
grown at 3 soil N levels in 1985.
( t a l l group)
Chlorophyll Concent.
E.
Dough
M.
Dough
H.
Dough
0.77
0.78
0.78
0.77
0.75
0.80
0.77
0.80
0.81
0.78
0.79
0.76
0.78
0.78
0.72
0.74
0.75
0.81
0.76
0.78
0.77
0.75
I .99
2.09
1.90
2.04
2.17
1.89
2.04
1.95
2.03
1.58
2.02
1.76
I .98
I .98
2.11
2.09
2.10
1.97
0.677
1.050
0.463
1.043
0.910
2.00
1.32
1.61
0.96
1.58
1.73
1.15
1.47
1.48
1.48
0.77
1.41
0.92
1,23
1.43
1.84
1.70
1.74
1.36
1.37
1.24
1.69
1.3,0
3.5
0.77
1.99
1.40
0.827
2.8
10.6
5-4
6.7
1.5
0.2
; 0.2
0.02
0.07
0.10
0.096
0.7
2.1
0.3
0.3
0.03
0.10
0.14
0.135
1.0
GY
GPP
TPP
NHI
Brule (USA,NE)
Grana (Poland)
Bezost.l (USSR)
AWl2399 (E.Germ.)
Arena ( S w i t z e r l . )
NE7060 (USA,NE)
Odissa-4 (USSR)
At las-66 (USA,IN)
Lavrin24 (Romania)
Lethb.32 (Canada)
Martonv.5 (Hungary)
Purdue (USA,IN)
Blueboy (USA,NC)
Houser (USA,NY)
Jana (Poland)
Alcedo (E.Germ.)
Aura (Finland)
MV-6 (Hungary)
Martonv. (Hungary)
NS2699 (Yugoslav.)
Redwin (USA,MT)
Lancota (USA,NE)
17.5
25.8
20.2
22.7
16.3
18.4
17,6
13.8
19.3
11.8
19.2
13.2
15.7
22.6
20.8
19.0
16.0
20.6
13.6
21.0
16.3
13.3
13.7
14.1
15.6
15.2
16.1
15.6
18.3
14.1
14.7
14.8
14.2
14.4
13.5
13.0
13.7
14.9
14.5
13.8
14.1
16.1
15.6
3-0
4.5
3.7
4. 5
3-3
3.7
3-6
3.2
3.4
2.2
3-6
2. 5
2.9
4.0
3.7
3.6
3.2
4.2
3.8
2. 4
4.4
3.4
Means
18.4
14.7
C.V.
14.4
LSD.05
LDS.01
Cu lti va rs
23.2
1.99
1.93
2.13
12.8
ChL.D.
0.663
0.753
1.050
0.803
0.190
0.583
0.523
0.817
0.767
1.383
1.223
1.233
O.683
0.713
0.650
1.280
0.743
21.0
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
Protein; NHI = Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
40. I
39.3
37.6
37-6
37.1
43.2
39-3
38.9
40.7
38.0
37.6
42.5
40.7
40.9
34.0
35.4
32.6
36.8
40.7
38.7
40.4
40.8
38.8
3.3
43
Table 15.
Eight t r a i t means of 22 w in te r wheat c u l t i v a r s (medium height
group) grown at 3 soil N levels in 1985.
Chlorophyll Concent.
TPP
NHI
E.
Dough
M.
Dough
H.
Dough
0.847
0.860
0.897
0.697
0.963
0.650
0.307
0.990
0.747
0.050
0.473
0.877
0.533
1.260
0.373
0.357
0.187
0.580
1.070
39.9
42.3
38.7
41.5
34.8
39.6
39.8
41.6
41.9
33.2
39.0,
38.7
38.3
37.1
41.5
41.3
34.5
38.3
40.2
38.4
40.0
40.0
0.693
39.1
ChL.D.
GY
GPP
13.2
13.2
18.5
14.2
21 .4
11.4
18.5
10.9
9-9
13.1
15.5
16.0
16.8
13.6
14.8
9.5
17.8
10.5
12.1
18.3
22.0
2. 6
13.3
4.0
13.5
14.0
2.3
3.5
14.5
2.5
13.7
13.5
3.9
2. 0
14.0
3.3
13-9
13.4
1.9
15.0
1.9
2.4
13-7
13.8
2.9
15,8
3.1
2.9
12.9
14.1
2.5
2.9
15.7
13.2 . 1.7
3.2
13.8
13.6
1.9
2.4
15.7
3.4
13.7
4. 6
15.1
0.75
0.78
0.81
0.76
0.77
0.75
0.78
0.78
0.76
O.78
0.76
0.74
0.83
0.76
0.77
0.82
0.75
0.77
0.78
0.78
0.74
0.72
2.13
1.89
2.01
1.95
1.89
1.31
2.02
2.04
1.19
1.81
2.05
1.98
2.08
1.95
1.95
1.82
1.75
1.80
1.51
1.84
2.03
1.46
1.42
1.29
1.29
1.02
1.25
0.67
1.53
1.48
0.38
1.19
1.44
I .22
1.69
1.33
1.48
1.64
1.03
0.78
0.51
1.03
1.50
15.2
14.2
2. 8
0.77
1.87
I .21
12.8
2.4
13.3
LSD.05
1.1
0.2
0.2
0.02
0.13
0.09
0.085
0.7’
LSD.Ol
1.5
0.3
0.3
0.03
0.18
0.13
0.120
1.0
Cult i v a r s
Sutjeska (Yugos.)
Lavrin-32 (Romania)
Orov. (Yugoslavia)
F29-75 (Romania)
MV-7 (Romania)
Horosh (Japan)
CA8055 (China)
NS-15-89A (Yugosl.)
K a t i a - I (Bulgaria)
Feng-Kang (China)
Saiente ( I t a l y )
F29-76 (Romania)
GK-Prot. (Hungary)
Clement ( N et he r ld .)
Doina (Romania)
WWP.4394 ( Austria)
Bastion ( N et h er ld .)
TX71A562-6 (USA1TX)
WW330 ( A u s t r a l ia )
Chokwang (Korea)
Centurk (USA1NE)
MT 7811 (USA1MT)
Means
C.V.
.
23.0
4.4
2.08
12.4
14.0
1.320
0.613
0.700
22.4
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
Protein; NHI - Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
3.3
Table 16.
Eight t r a i t means of 22 w in te r wheat c u l t i v a r s
grown at 3 soil N levels in 1985.
(short group)
Chlorophyll Concent.
E.
Dough
M.
Dough
H.
Dough
1.07
0.327
0.820
0.547
0.787
0.640
0.883
0.193
0.357
0.527
0.890
1.377
O.O83
0.680
0.170
0.400
ChL.D.
GY
GPP
TPP
NHI
Vratza (Bulgaria)
Daws (USA,WA)
MV-22-27 (Hungary)
Vala (Czechslov.)
Stephens (USA,OR)
Ogosta (Bulgaria)
WWP.4258 ( Austria)
Vega (Bulgaria)
NS2630-1 ( Yugosl.)
Sudova S. (Bulgar.)
Loudog. (Bulgaria)
Bounty (England)
Pai Yu P. (China)
NS2699 (Yugoslav.)
NE79Y90576 (USA,NE)
Adams (Au stria )
Inernio ( I t a l y )
Trakia (Bulgaria)
Marisml. (England)
PL V (USA,KA)
MT 7877 (USA,MT)
15.5
21.0
19.4
20.0
20.5
13-9
12.0
16. 6
13.5
21.9
13-9
11.1
16.6
11.0
12.6
25.3
12.1
12.7
9.5
11.2
21.8
14.0
13.3
13.4
13-2
13-7
14.4
14.2
14.5
13-7
14.3
13.9
15.8
13.6
14.6
14.0
14.7
13.6
13-6
14.2
16.1
14.3
2.8
3.6
3-5
3.6
3-5
2.6
2 .1
3.1
2.4
4.2
2.5
2-3
2.9
2.1
2.3
4. 6
2.1
2.2
1.9
2.3
4.4
0.79
0.77
0.74
0.73
0.79
0.80
0.80
0.78
0.80
0.75
0.77
0.78
0.79
0.78
0.77
0.81
0.81
0.79
0.69
0.77
0.71
1.91
1.99
1.93
1.82
2.05
I .96
1.61
1.88
1.90
1.79
1.95
1.22
I .99
1.53
1.81
2.00
1.96
1.99
1.54
1.97
0.37
1.32
0.78
1.22
1.62
1.06
1.17
1.71
0.71
1.43
Means
16.2
14.2
3-0
0.77
1.85
1.20
C.V.
12.6
1.9
11.5
3.4
7.4
LSD.05
1.1
0.2
0.2
0.01
0.08
0.07
0.078
0.7
LSD.Ol
1.6
0.2
0.3
0.02
0.11
0.12
0.111
1.0
Cu lti va rs
1.83
1.42
I .05
1.52
I .42
1.35
0.88
0.97
1.23
1.39
1.63
13.2
1.117
0.297
0.500
1.363
0.370
0.927
0.637
22.5
GY = Grain Y i e l d ; GPP = Grain Protein Percentage; TPP = Total Plant
Protein; NHI = Nitrogen Harvest Index; ChL.D. = Chlorophyll Duration
39.1
39.3
37.6
38.7
38.2
37.9
36.2
37-7
41.3
41.7
33.6
35.2
38.6
37.9
38.8
39-8
31.5
40.7
32.6
39.7
40.8
38.0
3.3
45
Table 17.
Soil
N Levels
T r a i t means of 22 t a l l c u l t i v a r s grown in 1984 and 1985 at 6
soil N le v e l s .
Chlorophyl I Concent.
Anth. E.D.
M.D.
H.D.
ChL.D.
2.45
2.36
1.15
0.27
44.2
0.779
2.50
1.78
I .04
0.22
42.1
8.8
0.763
2.52
1.57
1.57
0.20
40.0
0.2
0.9
0.028
0.05
0.15
0.14
0.08
0. 8
0.05
0.5
2.0
0.058
0.11
0.30
0.28
0.16
1.7
17-9
0.45
13.8
3.1
0.784
2.23
2.08
1.42
0.80
39.4
NI-(ISZ)
19-9
0.43
14.9
3.8
0.774
2.35
2.02
1.46
0.84
39.2
N2-(242)
17.3
0.43
15.5
3.5
0.756
2.17
1.86
1.31
0.77
37-8
LSD.05
0.7
0.05
0.2
0.1
0.014
0.06
0.06
0,05
0.06
0.3
LSD.01
1.5
0.10
0.3
0.2
0.029
0.12
0.12
0.10
O'. 1.3
0. 7
GY
HI
GPP
TPP
NHI
NO-(IlO)+
50.7
0.39
13.8
9.2
0.756
N I - ( 1 95)
49.8
0.41
13.9
8.9
N2-(280)
48.4
0.38
14.0
LSD.05
4.7
0.02
LSD.01
9-7
NO-(22)
1984
1985
GY = Grain Y i e l d ; HI = Harvest Index; GPP = Grain Protein Percentage;
TPP = Total Plant Protein; NHI = Nitrogen Harvest Index; E.D. = Early
Dough; M.D. = Medium Dough; HvD. = Hard Dough; ChL.D. = Chlorophyll
Duration
+ = Kg/ha Soil Nitrogen ( a v a i l a b l e + added)
Table 18.
Soil
N Levels
T r a i t means of 22 medium height cult!vans grown in 1984 and
1985 a t 6 soil N le v e l s .
Chlorophyl I Concent.
Anth. E.D.
M.D.
H.D.
ChL.D.
2.49
2.38
0.99
0.25
45.5
0.787
2.51
1.74
0.86
0.18
44.3
8.3
0.753
2.54
1.43
1.44
0.14
41.9
0.2
0.6
0.022
0.05
0.17
0.16
0.06
0. 8
0.05
0.5
1.3
0.045
0.10
0.36
0.33
0.13
1.6
13.6
0.44
13.3
2,3
0.774
2.22
2.00
1.33
0.58
39.4
NI - ( 132)
15.8
0.44
14.4
3.0
0.764
2.29
1.83
1.21
0.57
38.6
N2-(242)
16.1
0.44
14.8
3.1
0.766
2.16
1.77
1.15
0.61
39-3
LSD.05
0.3
0.01
0,1
0.1
0.01 I
0.04
0.13
0.05
0.07
0.3
LSD.01
0.5
0.02
0.2
0.2
0.023
0.08
0.27
0.11
0.14
0. 7
GY
HI
GPP
TPP
NHI
NO- (IlO)+
45.1
0.42
13-4
7-7
0.776
N I -(1 9 5)
46.7
0.43
13.7
8.0
N 2 -(280)
46.0
0.39
13.9
LSD.05
3.9
0.02
LSD.01
8.0
NO-(22)
1984 ;
1985
GY = Grain Y i e l d ; HI = Harvest Index; GPP = Grain Protein Percentage;
TPP = Total Plant Protein; NHI = Nitrogen Harvest Index; E.D. = Early
Dough; M.D. = Medium Dough; H.D. = Hard Dough; ChL.D. = Chlorophyll
Duration
+ = Kg/ha Soil Nitrogen ( a v a i l a b l e + added)
Table 19.
Soil
N Levels
T r a i t means of 22 short c u l t i v a r s grown in 1984 and 1985 at
6 soil N le v e l s .
Chlorophyll Concent.
Anth. E.D.
M.D.
H.D.
ChL.D.
2.50
2.38
I .01
0.22
45.2
0.800
2.58
1.71
0.93
0.17
42.9
7.8
0.779
2.55
1.39
1.38
0.14
41.4
0.2
0.8
0.025
0.04
0.08
0.09
0.04
0. 7
0.05
0.4
1.7
0.051
0.08
0.16
0.19
0.08
1.4
15.5
0.45
13.3
2.5
0.788
2.25
2.06
I .36
0.63
39.3
NI -(1 3 2 )
16.8
0.43
14.4
3.1
0.773
2.31
1.91
1.19
0.53
37-0
N2-(242)
16.2
0.44
15.1
3.2
0.756
2.18
1.57
1.04
0.38
37.8
LSD.05
0.6
0.01
0.1
0.1
0.013
0.04
0.06
0.05
0.03
0.3
LSD.01
1 .2
0.02
0,3
0.2
0.026
0.08
0.13
0.10
0.07
0. 7
GY
HI
GPP
TPP
NHI
NO- (IIO)+
47-1
0.44
13.1
7.7
0.800
NI -(1 9 5)
46.2
0.44
13.3
7.7
N 2 - (280)
45.7
0.41
13.6
LSD.05
4.6
0.03
LSD.01
9-6
NO-(22)
1984
1985
GY = Grain Y i e l d ; HI = Harvest Index; GPP = Grain Protein Percentage;
TPP = Total Plant Prot ein ; NHI = Nitrogen Harvest Index; E.D. = Early
Dough; M.D. = Medium Dough; H.D. = Hard Dough; ChL.D. = Chlorophyll
Duration
+ = Kg/ha Soil Nitrogen ( a v a i l a b l e + added)
MONTANA STATE UNIVERSITY LIBRARIES
stks N378.AI49
Grain protein and grain yield as functio
3 1762 00512055 3
"'"In
N37R
Afbg
cot ) .
2