Relationship between seed vigor testing and field performance of regar meadow bromegrass (bromus
biebersteinii Roem and Schult)
by Reginald Denny Hall, II
A thesis submitted in partial fulfillment of the requirements for the degree of Master of Science in
Agronomy
Montana State University
© Copyright by Reginald Denny Hall, II (1987)
Abstract:
The standard germination test continues to be the primary method for determining seed quality.
However, germination results often overestimate actual field emergence. Seed quality in grasses has
been shown to affect field emergence and subsequent stand establishment. Several physiological and
biochemical tests have been developed for evaluating seed vigor in various crops, but the relationship
between vigor testing and field performance has not been studied extensively in grasses.
This study evaluated several laboratory vigor tests to determine their relationship to field performance
of 'Regar' meadow bromegrass. Standard germination and accelerated aging tests were used to evaluate
four seed lots during preliminary studies in 1985. Field performance of these lots were evaluated by
determining speed of emergence, total emergence, and forage yield. Total emergence and forage yield
were significantly correlated with accelerated aging. The following year, standard germination,
seedling growth rate, accelerated aging, electrical conductivity, respiration rate, and ATP content were
used to evaluate ten seed lots planted in the field. Accelerated aging, respiration rate, and ATP content
were significantly correlated with forage yield. Multiple stepwise regression selected accelerated aging
and respiration rate as the best model for predicting first year forage yield. RELATIONSHIP BETWEEN SEED VIGOR TESTING AND FIELD
PERFORMANCE OF ,REGAR1 MEADOW BROMEGRASS
(Bromus biebersteinii Roem and Schult)
by
Reginald Denny Hall, II
A thesis submitted in partial fulfillment
of the requirements for the degree
of
Master of Science
in
Agronomy
MONTANA STATE UNIVERSITY
Bozeman, Montana
May 1987
M LI
L-Iw
MAIN
/l/37f ,
ii
APPROVAL
of a thesis submitted by
Reginald Denny Hall, II
This thesis has been read by each member of the thesis
committee and has been found to be satisfactory regarding
content, English usage, format, citations, bibliographic
style, and consistency, and is ready for submission to the
College of Graduate Studies.
Chairperson,Graduate Committee
Approved for the Major Department
Date
Head, Major Department
Approved for the College of Graduate Studies
Date
Graduate Dean
iii
STATEMENT OF PERMISSION TO USE
In presenting this thesis in partial fulfillment of the
requirements
for
a
master's
degree
at
Montana
State
University, I agree that the Library shall make it available
to
borrowers under rules of the Library.
from
Brief
this thesis are allowable without special
quotations
permission,
provided that accurate acknowledgment of source is made.
Permission for extensive quotation from or reproduction
of
this thesis may be granted by my major professor,
his
absence,
by
opinion of either,
scholarly
this
when,
in
the
the proposed use of the material is
for
purposes. Any copying or use of the material
in
thesis for financial gain shall not be allowed without
Signature.
Date.
the Director of Libraries
or in
V
ACKNOWLEDGMENTS
I
wish
to
express my
sincere
appreciation
to
the
guidance,
and
following:
Dr.
Loren
friendship
Wiesner for his assistance,
as my major professor.
Additional
appreciation
for acquainting me to grass seed production and accepting me
as his research technician for the past two years.
Dr.
Ray
Ditterline for his
advice,
friendship,
and
support while serving on my graduate committee.
John Scheetz,
Materials
Center
his staff,
for
and the S.C.S. Bridget Plant
assistance
in
field
research
and
providing support for my research position.
Dr.
J.
M.
Martin for his assistance with statistical
data analyses.
My
parents and sister for providing encouragement
and
support in pursuing my advanced degree.
To
my fellow graduate students for
their
friendship,
assistance, support, and good times shared together. Special
thanks to Larry S .
Hicks for providing an excellent fishing
guide service.
I
vi
TABLE OF CONTENTS
Page
APPROVAL.....................
STATEMENT OF PERMISSION TO USE.......................
VITA.......................
ACKNOWLEDGMENTS...............
TABLE OF CONTENTS.........
LIST OF TABLES.......................................
ABSTRACT.............. ............................ ..
ii
iii
iv
V
vi
viii
ix
Chapter
I.
, II.
I
LITERATURE REVIEW..........
3
CO ^ IT) f" o
III.
INTRODUCTION................................
Significance of Vigor Testing
Seedling Growth Rate........
Accelerated Aging...........
Electrical Conductivity.....
Respiration................ .
Adenosine Triphosphate Content...........
11
RELATIONSHIP BETWEEN SEED VIGOR TESTING
AND FIELD PERFORMANCE OF 'REGAR1 MEADOW
BROMEGRASS (Bromus biebersteinii Roem and
Schult)............................
14
Introduction.........
Materials and Methods......................
Laboratory Studies.............
Standard Germination Test...........
Seedling Growth Rate................
Accelerated Aging.................
Electrical Conductivity...............
Re S p i r a t l O n . . . . . . . . . . . . . . . . . . . o o e e e e . e
ATP Content...................
14
15
16
16
16
17
18
I9
19
vii
TABLE OF CONTENTS-Continued
Page
Field Studies...........................
Statistical Analysis...........
Results and Discussion.........
LITERATURE CITED..................................... .
20
22
22
32
I
APPENDIX.........
40
viii
LIST OF TABLES
Table
1.
2.
3.
4.
5.
6.
7.
Page
Mean comparisons among standard germination,
accelerated aging, and field performance
variables for four seed lots of Regar meadow
bromegrass - Bozeman, MT. 1985.................
23
Mean comparisons among standard germination,
vigor tests, and field performance variables
for 10 seed lots of Regar meadow bromegrassBozeman, MT. 1986............. .................
25
Simple correlations among standard germination,
accelerated aging, and field performance
variables for four seed lots of Regar meadow
bromegrass - Bozeman, MT. 1985..............
26
Simple correlations among standard germination,
vigor tests, and field performance variables
for 10 seed lots of Regar meadow bromegrass Bozeman,
Mt. 1986.............
27
Summary of multiple stepwise regression comparing
first year forage yield with vigor tests for 10
seed lots of Regar meadow bromegrass - Bozeman,
MT. 1986.......
31
Simple correlations among standard germination,
accelerated aging and field performance variables
for four seed lots of Regar meadow bromegrass Bozeman,
MT. 1985..........................
41
Simple correlations among standard germination,
vigor tests, and field performance variables
for 10 seed lots of Regar meadow bromegrass Bozeman, MT. 1986......
42
ix
ABSTRACT
The standard germination test continues to be the
primary method for determining seed quality.
However,
germination
results
often overestimate
actual
field
emergence. Seed quality in grasses has been shown to affect
field emergence and subsequent stand establishment. Several
physiological and biochemical tests have been developed for
evaluating seed vigor in various crops, but the relationship
between vigor testing and field performance has not been
studied extensively in grasses.
This study evaluated several laboratory vigor tests to
determine their relationship to field performance of 'Regar1
meadow bromegrass. Standard germination and accelerated
aging tests were used to evaluate four seed lots during
preliminary studies in 1985. Field performance of these lots
were evaluated by determining speed of emergence, total
emergence, and forage yield. Total emergence and forage
yield were significantly correlated with accelerated aging.
The following year, standard germination, seedling growth
rate,
accelerated
aging,
electrical
conductivity,
respiration rate, and ATP content were used to evaluate ten
seed
lots planted in the field.
Accelerated
aging,
respiration
rate,
and ATP content were significantly
correlated with forage yield. Multiple stepwise regression
selected accelerated aging and respiration rate as the best
model for predicting first year forage yield.
I
CHAPTER I
INTRODUCTION
lRegar1 meadow
and
bromegrass (Bromus biebersteinii
Schult), an improved cool season grass
selected
Roem
cultivar,
for its rapid regrowth characteristics.
Regar
was
is
winterhardy, moderately drought tolerant/ and its bunchgrass
habit
is
compatible
grasses and legumes.
and
seed
production
when
planted
with
other
perennial
Regar is a popular cultivar for forage
in the
northern
United
States
and
Canada.
Stand
species7
establishment
including
encountered. Adverse
cultural
practices
establishment.
These
problems of small
Regar
meadow
seeded
bromegrass,
is
environmental conditions or
can
result
in
forage
improper
erratic
factors impose stress on
often
stand
germinating
seeds and growing seedlings, depleting their energy supply/
and subsequently
is
preventing their emergence.
Therefore, it
important to plant high quality seed which possess
potential
to germinate and produce rapid growing
the
seedlings
under a wide range of field conditions.
The standard germination test is the primary method for
determining seed quality. However, germination results often
overestimate
actual field emergence.
Germination tests are
2
performed under optimum conditions seldom encountered in the
field.
The seed vigor concept was developed to provide a
accurate
appraisal
of seed quality as it relates to
emergence . . Laboratory
physiological
and
seed
biochemical
vigor
tests
methods
to
more
field
utilize
both
evaluate
seed
emergence
and
vigor in various crops.
Grass
seed
quality
affects
field
subsequent stand establishment. However, the relationship of
vigor
tests
with
field performance has not
extensively in grasses.
evaluate
been
studied
The objective of this study was
to
several laboratory vigor tests and determine their
relationship
bromegrass.
to
field
performance
of
Regar
meadow
3
CHAPTER II
LITERATURE REVIEW
Significance of Vigor Testing
The
"Rules
Association
of
Official
Seed
Analysts
(AOSA),
for Testing Seeds" (1981) defines germination in the
laboratory as,
"the emergence and development from the seed
embryo of those essential structures which, for the kinds of
seed in question, are indicative of the ability to produce a
normal
plant,
conditions,
germination
under favorable conditions." Whenever
at
planting,
tests
correlate
(Abdalla and Roberts,1969;
1979).
However,
optimum.
Under
germination
are
near
well
Perry,
optimum,
with
1977;
field
field
standard
emergence
Egli and Tekrony,
actual field conditions are often far from
suboptimum
field
conditions,
standard
tests differ significantly from field emergence
for a given seed lot (Sherf,
1953;
Delouche and
Caldwell,
1960; Tekrony and Egli, 1977; Johnson and Wax, 1978; Yaklich
and Kulik, 1979; Naylor, 1981).
The seed vigor concept was developed to provide a
accurate
appraisal of seed quality in relationship to field
emergence.
The
extensively
McDonald
more
by
definition of seed vigor has been
Heydecker
(1975,1980a).
(1972),
The
Woodstock
International
reviewed
(1973),
Seed
and
Testing
4
Association (ISTA),
properties
and
defined seed vigor as the "sum of those
which determine the potential level of
performance
of
the seed lot
seedling emergence" (Perry,
by
during
1978).
germination
Seed vigor was
AOSA as "those properties which determine the
for
rapid,
uniform
emergence,
activity
and
defined
potential
and development of
normal
seedlings under a wide range of field conditions" (McDonald,
1980b).
Seed vigor testing has become an increasingly important
component
of seed
analysis
(McDonald,
1975).
Successful
vigor tests are reproducible, correlate with emergence under
field conditions, rapid, objective, simple, and economically
practical
(AOSA,
differentiate
under
field
Naylor
1983).
Vigor
tests
are
designed
to
between seed lots of high and low performance
conditions
(Heydecker,
1972).
Marshall
(1984) reported that improved grass emergence
be achieved by planting high vigor seed;
and
could
that is, seed that
establishes well under poor conditions.
Seedling Growth Rate
Seedling
standard
content
growth rate tests are conducted similarly
germination test?,
of media.
but require specific
to
moisture
Seedling growth is measured in terms
of
dry weight or linear growth after a given period of growth.
'I
Woodstock (1969a) reported that measuring linear growth
5
of corn (Zea mays L .) seedlings was a sensitive indicator of
seed quality. Even under optimum conditions, seedling growth
rate
tests
revealed
vigor
differences
not
detected
by
percentage germination (Woodstock and Grabe, 1967).
Perry
(1977)
evaluated plumule growth as
vigor test for barley (Hordeum vulgare L.).
and
than
conditions
the
were
correlations
standard
germination
unfavorable.
were
found
between shoot
with
test
However,
possible
Field emergence
grain yield were more closely correlated
length
a
plumule
when
no
soil
significant
length
and
field
emergence for soybeans (Glycine max L . Merr.) (Yaklich
and
Kulik, 1979).
Seedling
dry
weight
has been suggested by
Vigor Testing Committee (Woodstock,
of
soybean seedlings,
associated
AOSA
1976). Total dry weight
excluding cotyledons,
with vigor (Edje and
the
Burris,
were
1970).
closely
Mock
and
McNeill (1979) evaluated cold tolerance responses in several
inbred
after
lines of corn.
planting
Seedling dry weight sampled 42
correlated significantly with grain
days
yield.
Accelerated Aging
The
accelerated
germination.
C)
and
depending
high
on
aging
test stresses
seed
prior
to
Seeds are subjected to high temperature (40-45
humidity (nearly
the kind of seed,
100%)
for
varying
after which a
times,
germination
6
test
is
performed.
tolerate
the
betterr
theory,
high
vigor
high temperature - high
seed
humidity
and retain its ability to germinate.
developed
to
(Delouche,
suggested
which
In
predict
using
seed
the
will
of
well
seed
1973).
accelerated aging test
perform
conditions
This test was
relative storability
Delouche and Baskin,
1965;
under
should
lots
Later, AOSA
to
determine
field
conditions
(Woodstock, 1976) .
The first method involved placing seed in open
boxes
and then inserting the boxes inside a
where
humidity and temperature were held
(1977)
modified
the
plastic
large
chamber
constant.
procedure by placing
seeds
Baskin
in
wire
baskets over a measured amount of water in a sealed jar, and
placing
the jars in a chamber. McDonald (1978) found
that
the
original seed moisture of soybeans and barley influence
the
degree
test.
of deterioration during the
Subsequently,
McDonald
and
accelerated
Phaneedranath
aging
(1978)
further modified the accelerated aging test by placing seeds
in a single layer on wire mesh trays over a measured
amount
of
(1979)
water
reported
method
inside
that
plastic
the
results
in
germination
addition of 40 ml.
minimum
test
boxes.
Tao
water to
variation.
the
This
tray
method
produced, excellent repeatability in a national vigor referee
test
for soybeans (Tao,
Subcommittee
suggested
1980).
In 1984,
that McDonald
and
the
AOSA
Vigor
Phaneedranath1s
7
(1978)
method
be
adopted as the
recommended
method
for
accelerated aging (Tekronyf 1985).
Accelerated
performance
aging
of
bean
has
been
(Phaseolus
used
to
predict
vulgaris
L.)
field
(Roos
and
Manalof 1971), soybean (Byrd and Delouchef 1971; Tekrony and
Eglif 1977; Kulik and Yaklich, 1982)f and
hirsutum
L.)
(Bishnoi
and
cotton (Gossypium
Delouchef
1980) .
Controlled
deterioration, a technique similar to accelerated agingf has
been
significantly
Italian
ryegrass
correlated
(Lolium
with
field
multiflorum
L.)
emergence
(Marshall
of
and
Naylor , 1985) .
Electrical Conductivity
The
leakage
electrical conductivity test measures
from
deteriorating seeds.
Ching
and
electrolyte
Schoolcraft
(1968) reported that deteriorated seeds leach more
cellular
components than nondeteriorated seeds , presumably because of
increased
membrane permeability.
Roberts (1979) associated
the loss of seed viability with a loss of membrane integrity
resulting in the enhanced efflux of cellular constituents in
seed leachates.
Membrane
reactions
that occur in living cells.
ultrastructure
aestivum
integrity is important for
L.)
and
and
permeability
many
biochemical
Changes in
in aged
pea (Pisum sativum L.)
wheat
seed
membrane
(Triticum
have
been
8
detected
by
electron
microscope
studies
(Anderson
et
al.,1970; Harman and Granett, 1972).
Abdul-Baki
and
Anderson (1970) reported
that
leaching from barley seed appears to be regulated
by
sugar utilization rate during germination,
changes
in
dry
seed
membrane
sugars
primarily
rather
permeability.
than
Glucose
utilization rate was faster in high quality seeds,
and
the
glucose concentration in the leachate was lower.
Woodstock et al. (1985) suggested that mineral leaching
in
weathered
cotton seeds was a better indicator
of
seed
vigor than the total release of electrolytes. The release of
+
++
K
and Ca
was significantly correlated with measurements
of seed quality.
Electrical conductivity measurements were significantly
correlated
AOSA
with field emergence in corn and soybean
refereed vigor test
(Tao,
1980a;
Tao,
in
1980b).
an
The
results were also repeatable among seed laboratories.
The
electrical conductivity test correlates
vigor in seeds of pea (Matthews and Bradnock,
well with
1967;
Carver
and Matthews, 1975), rice (Oryza sativa L .) (Agrawal, 1977),
corn (Gill and Delouche,
soybean
1979;
Tao,
(Abdul-Baki and Anderson,
Tao,
1980a;
Tao,
incarnatum L. 'Dixie')
(Shorea
1973;
1980a;
1973;
Tao,
Yaklich
1980b) ,
et
al.,
1980b), crimson clover (Trifolium
(Ching and Schoolcraft, 1968) and sal
robusta Gaertn.
f .) (Nautiyal and Purohit,
1985).
9
However, poor correlations between electrolytic leakage and
seed vigor have been reported in sorghum (Sorghum bicolor L.
Moench)
(Perl and Gelmondf
1978)f
muskmelon (Cucumis
meIo
L .) (Pesis and Ngf 1983)f and Italian ryegrass (Marshall and
Naylorf 1985).
Respiration
Seed
germination and seedling growth require metabolic
energy supplied by respiration. Vigorous seed germinates and
grows
more rapidly than nonvigorous
energy
through
(1975)
reported
respiratory
increased
seed,
respiratory
that mitochondrial
requiring
activity.
more
McDonald
dysfunction
decreases
activity and results in little or no
embryonic
axis elongation.
Vigor evaluation by respiration tests are quantitativef
rapidf
easy
routinely
to standardize and perform,
testing,
precautions,
many seed
with
treatments can influence seed respiration. Pretreating
corn
temperatures
bean
(Phaseolus
reduced
1966).
suitable
seed
lima
(Woodstock,
and,
However,
and
reliable
samples
well - suited for
lunatus
respiration and
growth (Woodstock and Feeley,
1965;
L.)
with
subsequent
chilling
seedling
Woodstock and Pollock,
1965), respectively. Exposing corn seed to gamma - radiation
treatments
markedly
inhibited
seedling
respiration rate (Woodstock and Combs,
1965).
growth
and
Mechanically
10
injured seed was less vigorous and had increased rather than
decreased respiration rates (Woodstock,
of
1969b). Respiration
pea embryo axes infected with Aspergillus ruber Thom and
Church
was
lower than noninfected
peas
after
imbibition
(Harman and Drury, 1973).
Bewley
and Black (1982) reported respiratory
patterns
exhibited by deteriorating seeds are variable, and sometimes
precede,
Bewley
accompany,
or lag behind germination.
(1984) suggested that the correlation between oxygen
uptake and seedling vigor varies with time after
at
Halmer and
which
respiration is measured,
imbibition
and the number of
days
after which seedling vigor is determined.
Respiration
performance
DasGupta
1969;
correlates
in
and
seeds
Matthews,
of wheat
Austenson,
Abdul-Baki
well
(Kittock
1973) ,
and Anderson,
1975) ,
with
vigor
or
field
and
Law,
1968;
soybeans (Burris et
1973),
peas
al.,
(Carver
and
corn (Woodstock and Grabe, 1967; Cantrell
et al., 1972), cotton (Bishnoi and Delouche, 1980; Woodstock
et
al.,
1985) ,
and Italian ryegrass (Marshall and Naylor,
1985).
However,
some
researchers
have
correlations
between respiration and seed
Baki
observed
(1969)
decrease
in
oxygen
that
uptake
in
reported
quality.
deteriorating
lagged
behind
poor
Abdul-
barley
the
changes
in
germination, shoot growth, and glucose utilization. Anderson
11
(1970)
reported
correlate
although
older
with
that
in barley,
either
oxygen
percentage
uptake
did
germination
not
or
age,
respiratory quotients were consistently higher
seeds.
Byrd
deteriorating
behind
and Delduche (1971)
observed
that
in
in
soybean the decrease in oxygen uptake ■lagged
germination
however, respiratory
and
seedling
quotients
increased
growth
as
rate;
germination
decreased. Bonner (1974) reported that germination rates did
not correlate with oxygen uptake in deteriorating cherrybark
oak (Ouercus falcata var. pagodaefolia) acorn.
Adenosine Triphosphate (ATP) Content
Seed
germination is an energy requiring
process.
energy
needed for biochemical reactions in living cells
stored
in high energy compounds such as ATP.
The
is
Extraction of
ATP has been performed in boiling water (Ching, 1973), or by
homogenizing
seeds
(Yaklich et al.,
luminescence
in
chilled
perchloric
acid
solution
1979). Extracted ATP is then measured by a
photometer
(Ching,
1973) , or
a
liquid
scintillation counter (Tao et al., 1974) using a luciferin luciferase
specialized
system.
This
method
is
rapid,
but
requires
equipment and trained personnel to conduct
the
test (AOSA, 1983).
Adenosine
seed imbibition;
triphosphate
is produced rapidly
following
however, the initial sources of ATP during
12
early
that
imbibition is still unknown.
the
Ching
(1972)
rapid ATP increase in seeds following
reported
imbibition
comes not only from oxidative phosphorylation, but also from
substrate
level phosphorylation such as
glycolytic
Willson
enzymes
and
are
Bonner
mitochondria
were
active in
(1971)
lacking
since
or
imbibed
seed.
that
peanut
seed
dry
found
in
glycolysis,
cytochrome
c
until
approximately 16 hours after imbibition started. Respiratory
activity
before that time was "alternate" respiration,
not sensitive to cyanide.
ATP
Moreland et al.
(1974)
and
observed
production in imbibed radish (Raphanus sativus L.) seed
under anaerobic conditions and in the presence of glycolysis
inhibitors.
They
reported
monophosphate
(AMP)
concentrations rather
diphosphate
(ADP)
decreases
during the early stage
in
adenosine
than
adenosine
of
germination,
suggesting an unknown system that phosphorylates AMP to ATP.
Perl
(1980) ,
developed
an
using
onion
(Allium cepa L .)
in vitro system for ATP
seed
synthesis
powder,
requiring
AMP, phosphoenolpyruvate (PEP), and orthophosphate.
Mitochondria isolated from seedling axes of new and old
soybean
efficiency
seed
when
exhibited
identical
added (Abu-Shakra and Ching,
differential
cofactors and
phosphorylative
substrates
1967) . The amount of inorganic
phosphate esterified into ATP per volume of oxygen
in
were
consumed
aged seeds was less than 50% that of unaged seeds.
They
13
suggested that mitochondria in aged seeds may be
uncoupled,
and
their inability to produce ATP
partially
efficiently
contributes to reduced germination vigor.
Ribonucleic
excised
from
acid
(RNA) and protein synthesis in
dry soybean seeds were
assayed
axes
after
being
subjected to varying accelerated aging treatments (Anderson,
1977) .
Lower
deteriorated
rates
of
RNA
and
protein
synthesis
seeds were associated with reduced ATP
in
tissue
content.
Adenosine
significantly
triphosphate
correlated
content in imbibed
with seedling weight
(Lactuca sativa L .) (Ching and Danielson,
crimson clover,
field
of
1972) ,
was
lettuce
ryegrass,
and common rape (Brassica napus L .) (Ching,
1973) . In soybean, ATP content
with
seeds
was significantly correlated
emergence in one trial , but
(Yaklich et al.,
not
in
another
1979). Pea seeds infected with Aspergillus
ruber Thom and Church exhibited decreased seed vigor and ATP
content
compared
However,
ATP
to uninfected peas (Tao
content
was
not
et
correlated
al.,
with
1974).
reduced
germination or vigor in corn, cucumber (Cucumis sativus L.),
onion, and radish seeds (Styer et al., 1980).
'14
CHAPTER III
RELATIONSHIP
BETWEEN
SEED
VIGOR TESTING
AND
FIELD
PERFORMANCE
OF
'REGAR1
MEADOW
BROMEGRASS
(Bromus biebersteinii Roem and Schult)
Introduction
The standard germination test is the primary method for
determining
seed
quality.
are
correlate
well with field emergence (Abdalla
Perry,
suboptimum
differ
optimum,
conditions,
field
planting,
1969;
near
When
standard
germination
and
at
tests
Roberts,
1977; Egli and Tekrony, 1979) „ However, under
field
conditions,
standard
germination
tests
significantly from field emergence for a given
lot (Tekrony and Egli,
seed
1977; Johnson and Wax, 1978; Yaklich
and Kulik, 1979; Naylor, 1981).
Seed
quality
in grasses affects field
emergence
subsequent stand establishment (Marshall and Naylor,
Several
physiological
developed
for
However,
these
extensively
respiration
and
evaluating
in
biochemical
seed
vigor
relationships have
grasses.
in
have
various
not . been
Controlled
tests have both been
tests
1984).
been
crops.
studied
deterioration
significantly
and
and
correlated
with field emergence of Italian ryegrass (Lolium multiflorum
L .)
(Marshall and Naylor,
correlates
well
with
1985) .
Electrical
germination
of
conductivity
crimson
clover
15
(Trifolium
incarnatum
L.
'Dixie'),
but
not
perennial
ryegrass (Lolium perenne L .) (Ching and Schoolcraft,
Poor
correlations
leakage
and
were also observed between
and field emergence of Italian
Naylor,
1985).
correlations
Ching,
1968).
electrolytic
ryegrass
(Marshall
(1973) observed significant
between ATP content in imbibed
ryegrass
seed
and seed weight or seedling size and weight.
Burris
measure
;
et
both
combinations
al. (1969)
questioned whether one test can
viability and
of
vigor.
Subsequently,
physiological and biochemical
various
tests
been utilized to predict field performance in several
have
crops
(Tekrony and Egli, 1977; Ram, 1983).
The
objective of this research was to evaluate
laboratory
vigor
several
tests to determine their relationship . to
field performance of Regar meadow bromegrass.
Materials and Methods
Seed
samples
State
lots evaluated were obtained from certified
seed
of Regar meadow bromegrass submitted to the Montana
University Seed Testing Laboratory.
Selections
were
based on varying levels of percentage germination. Four seed
lots
and
lower
were evaluated during preliminary studies in
one
lot (854) was. subjected, to
accelerated
quality prior to field planting.
evaluated during 1985-86.
1984-85,
aging
Ten seed lots
Two of these lots
to
were
(868 and 8610)
16
exhibited
and
one
low viability due to over-heating after
lot
(869) had been
harvesting and/or cleaning.
in
mechanically
at
injured
during
None of the seed lots evaluated
1985 were used in the 1986 studies.
stored
harvest,
All seed lots
room temperature (22C) in the laboratory
were
during
the course of these experiments.
Laboratory Studies
Standard Germination Test
Four groups of 100 seeds from each seed lot were placed
in
13
x 13.5 cm plastic germination boxes
moistened blotter papers.
containing
two
Seeds were germinated for 14 days
at alternating temperatures of ISC for sixteen hours and 2SC
for
eight hours (15-25C).
according
to
Official
Seed
"Rules
for
Analysts
Normal seedlings were
Testing
[AOSA],
Seed"
1981)
evaluated
(Association
and
expressed
of
as
percentage germination.
Seedling Growth Rate
Seedling growth rate test was conducted as suggested by
AOSA
(1983) with minor procedural modification.
procedure
was
replications.
The
basic
a rolled - towel germination test with
Each
roll consisted of two heavy weight
four
(#
76) paper towels, 25.4 x 38.1 cm, one below the seed and one
above the seed.
Towels were moistened with 30 ml water
per
17
towel.
Fifty
seeds per towel were arranged as described by
AOSA. The two paper towels Were loosely rolled within a 30.4
x 45.7 cm wax paper sheet and placed upright in a 15.5 x
cm container. Each container was
and
17
covered with a plastic bag
placed in a dark 15-25C germinator for 14 days.
Normal
seedlings were cut free from their caryopses and placed in 8
x
9 cm coin envelopes for drying.
Seedlings were dried
at
105C for 24 hours, then weighed to the nearest mg. Total dry
weight
was
determine
divided by the number of
normal
seedlings
to
seedling growth rate (mg/seedling).
Accelerated Aging
Preliminary accelerated aging tests during 1984-85 were
conducted
(1978)
as
recommended
with
seeds
were
covered
minor
evenly
by
procedural
McDonald
and
Phaneedranath
modifications.
distributed on wire mesh
Two-hundred
trays
plastic boxes containing 30 ml of water
and
inside
boxes
were placed in a Stults Scientific accelerated aging chamber
for eight days.
Temperature and humidity were maintained at
41C and 100% relative humidity,
aging
period,
seed
was
respectively. Following the
removed and standard
germination
tests conducted using four replications of 50 seeds per seed
lot.
Heavy
infestations
of
storage fungi
on
some
poor
quality seed lots made repeatability difficult.
During '1985-86 studies, the accelerated aging test was
18
conducted
as recommended by AOSA (Tekrony,1985) with
procedural
surface
modifications.
sterilized
minor
Four grams of seed per lot
in 30 ml of
1.5%
sodium
were
hypochlorite
(NaOCl) for 15 minutes, rinsed six times with sterile water,
oven-dried
at 32C for 24 hours,
and placed in a desiccator
24 hours prior to conducting accelerated aging
was uniformly
Plastic boxes were
inside a Stults Scientific accelerated aging chamber
for four days.
Temperature and humidity were maintained
41C and 100% relative humidity,
aging
Seed
distributed on wire mesh trays inside covered
plastic boxes containing 40 ml of water.
placed
tests.
period,
germination
seed
samples
tests conducted,
at
respectively. Following the
were
removed
and
standard
using four replications of 50
seeds per seed lot. An additional 50 seeds per seed lot were
removed,
weighed immediately to the nearest mg,
at
for
10SC
24 hours, and
re-weighed to
oven-dried
determine
seed
moisture content (wet - weight basis).
Electrical Conductivity
The
electrical
recommended
by
modifications.
lot
conductivity
AOSA
Four
(1981)
test
with
was
conducted
minor
as
procedural
replicates of 400 seeds from each seed
were weighed and placed in 50 ml test tubes
containing
40 ml deionized water. Tests tubes were incubated at 2OC for
24
hours,
after which time the contents of each test
tube
19
was
gently stirred, and
immediately.
Electrical
electrical
conductivity
conductivity
measured
was measured with
Markson
4503 conductivity meter and as umhos per gram
-I
weight (umhos g ).
a
seed
Respiration
,
Oxygen uptake was measured with
a Gilson
Differential
Respirometer. Four replicates of 25 seeds from each seed lot
were
imbibed
in
a
reaction
flask
containing
2
ml
of
distilled water for .24 hours, after which time 0.2 ml of 10%
KOH was added to the center well of each reaction flask. The
reaction
flasks
submerged
placed
on
the
respirometer
and
in
a 25C water bath and were shaken
-I
min . The system was'equilibrated for
oscillations
minutes
were
and readings were taken every hour for four
Respiration
rate
was
reported as
microliters
— I
absorbed per seed per hour (uLO
seed
of
78
30
hours.
oxygen
— I
hr
).
2
ATP Content
Four
were
replications of 0.5 g of seed from each seed
weighed
between
and
counted.
Each
two layers of Whatman No.
replication
was
lot
placed
I filter paper (9.0
cm)
which was moistened with 2.5 ml distilled water in a plastic
germination
box for four hours at 2OC.
Imbibed seeds
ground in a Wiley laboratory grinder (20 mm mesh) and a
were
0.3
20
g
sample removed and placed in a test tube for
Nine ml of dimethyl sulfoxide (99%) was
sample
and
mixed
extraction.
added to the ground
with a Vortex laboratory mixer
for
one
minute , after which time the particulate matter was allowed
to
settle
for two minutes.
transferred from each test
A 50
microliter
aliquot
was
tube into small plastic ependorf
tubes , and immediately diluted with 200 microliters of Hepes
buffer (pH 7.5) and placed on ice.
Extracted
quickly
Four replications of 150
transported for assaying.
microliters
TD-20e
Luminometer.
luciferin-luciferase
into
for
each
were
of the diluted seed extract were delivered into
small plastic test tubes ( 8 x 5 0 cm),
Turner
samples
and tubes placed in a
One - hundred
microliters
then
injected
test tube and the luminescence reading
recorded
each sample.
enzyme preparation was
of
Sample readings were compared to
an
ATP
standard curve and ATP content of each sample calculated as
-12
picograms (10
) of ATP per seed.
Dimethyl
sulfoxide (DMSO) used for ATP extraction
purchased from Sigma Chemical.
and
ATP standard,
was
Hepes buffer,
the Iuciferin-luciferase preparation was purchased from
Turner Designs.
Field Studies
All
at
the
seed lots evaluated in the laboratory were planted
Arthur
H . Post
field
research
laboratory
near
21
Bozeman, Montana.
variant
of
The
soil
is
classified
silt loam (fine-silty,
as
mixed family
Amsterdam
of
Typic
Haploborolls).
Four seed lots were planted in 1985 and 10 seed lots in
1986.
All
field experiments were conducted as a randomized
complete block with four replications.
x
1.83
Plot size was 6.09 m
m and each plot contained six rows spaced
apart.
Seeding
rate
0.305
was 15 pure live seed per 0.305 m
m
of
row, and seed was planted 12.7 mm deep.
After
0.5
m
planting and before emergence of seedlings,
of row were marked in
plot.
Emergence
obtained
calculated
described
from
rate
these areas.
similarly
by
and
Maguire
to
the two center rows
total
emergence
of
each
counts
were
Speed of emergence
speed
(1962) .
of
germination
Emergence
rate
two
index
index
index
was
as
was
calculated as follows:
X
=
number of seedlings
emerged
___________;
_________
increase of
seedlings emerged
from previous count
+ . . . + _____________________
days to first count
days to final
count
Forage yield was determined by harvesting 6.09 m of the
two center rows using a Rem flail harvester. Random moisture
samples were taken from each plot;
and
percentage
samples were oven-dried,
moisture determined on a wet-weight
basis.
22
Forage yield was corrected for moisture content and reported
-I
as kg ha
(dry-weight basis).
Statistical Analysis
All
Means
variables were subjected to analyses of
were
correlations
separated
were
using
computed
Neuman
-
the
mean
from
variance.
Kuels.
Simple
values
using
"MSUSTAT" (Lund, 1983) . A multiple stepwise regression model
building
vigor
procedure
tests
best
was used to determine which
predict
first year
forage
1986
seed
yield
(SAS
Institute Inc.,1985).
Results and Discussion
Significant
differences were found among seed lots for
•I
total seedlings emerged and forage yield in 1985 preliminary
studies
(Table
seedling
emergence
significantly
Speed
of
although
I).
higher
Seed
and
lot 852 had
first year forage
than
seed lot 854 for
emergence ratings did not
seed
the
differ
highest
yield
both
total
and
was
traits.
significantly,
lots which had high forage yields
also
had
high speed of emergence values.
Similar
field
results were obtained in 1986
seed lots were evaluated (Table 2).
were
when ten
Significant differences
found among seed lots for total seedling emergence and
first year forage yield. Seed lot 863 had the highest forage
23
yield and was significantly different from seven other
lots.
However,
seed
seed lot 864 had the highest total seedling
emergence and was significantly different than seed lots 869
and
868.
Speed
of
emergence
ratings
did
not
differ
significantly.
High variability among seed lots for speed of emergence
was
indicated by high coefficient of variation
percentages
of 69% (Table I) and 37% (Table 2) respectively for 1985 and
1986 studies.
seeding
depth
Small sample size (I meter of row) and uneven
and placement may have contributed
to
high
coefficient of variation for speed of emergence.
Table
I.
Mean comparisons among standard germination,
accelerated
aging,
and
field
performance
variables for four seed lots of Regar meadow
bromegrass - Bozeman, MT. 1985
Field parameters
Seed
Lot
Speed
of emerg.
Total
emerg.
Forage
yield
Std.
germ.
Accel.
aging
kg/ha
%
%
8.24
5.77
3.63
2.24
plants/m
I/
39b
, 36b
27ab
17a
1669b
1534ab
997 a
918a
88c
94d
84b
60a
33c
38c
20b
Oa
69
27
26
3
28
index
852
851
853
854
Laboratory parameters
2/
C.V.%
I/ Means followed by the same letter are not significantly
different
(P=O.05) according to Neuman-Keuls
mean
separation test
2/ Coefficient of variability
24
Laboratory
germination
were
I).
tests conducted during 1985
and accelerated aging.
were
standard
Significant differences
found among seed lots for both of these
tests
(Table
Seed lots 851 and 852, which had the highest first year
forage
yield
highest
and total seedling emergence,
standard
germination" percentage
also
and
had
the
accelerated
aging values.
Laboratory
germination,
tests
conducted
seedling
electrical
growth
conductivity,
Significant
in
1986
rate,
were
standard
accelerated
respiration,
and
ATP
aging,
content.
differences were found among seed lots for
laboratory tests (Table 2);
however,
all
some laboratory tests
were better predictors of field performance than others.
Standard
evaluated
during
commercially
standard
same
germination
acceptable
of
for
both years (Tables I and 2)
germination).
number
percentages
live
to
unacceptable
seed
lots
ranged
from
than
80%
Each seed lot was planted with
the
seed per meter
of
(less
row,
thus
seed
viability should not have influenced field performance.
Simple
tests
correlations were calculated
and field performance in 1985.
among
laboratory
Accelerated aging was
significantly correlated with total seedling emergence (r
0.97**)
and
first year forage yield (r = 0.88*),
with speed of emergence (Table 3).
correlated
but
=
not
Standard germination was
with total seedling emergence (r =
0.91*),
but
Table 2.
Mean comparisons among standard germination, vigor tests, and field performance variables
for ten seed lots of Regar meadow bromegrass - Bozeman, MT, 1986.
F ie ld parameters
Speed
T o ta l
o f emerg.
emerg.
Seed
Lot
Forage
y ie ld
S td .
germ.
kg Zha
%
index
plants/m
863
21.61
SSabiz
5582d
94ed
867
22.22
33ab
5245dc
866
34.31
47ab
868
20.31
864
SGR2z
mgZp la n t
Laboratory parameters
A ccel.
3Z
ECT
aging
2 8 4 .09d
43cb
. 79cbd
4.44a
242.35cb
69fe
.90cd
4.77a
2 . 12bcd
228.19b
51cd
.67cbd
3.34a
92ed
1.90ab
3 2 2 .16e
64fe
.55b
2.97a
4l49ab
90ed
1.85a
271.33cd
35b
. .52b
3.90a
4090ab
96e
2.20d
2 6 5 .72cbd
33b
. 64cb
4.01a
58ed
• 87cd
4.45a
15a
.79cbd
5.08a
13a
. 26a
3.49a
16
19
1.96abc
4986dc
88ed
2 . 22d
30a
4538bc
75c
38.42
55b
4207ab
865
30.31
38ab
861
19.40
32ab
862
22.50
35ab
3934ab
94ed
I . 96abc
249.09cbd
869
21.33
28a
3837ab
60b
I . 94abc
3 4 3 .61e
8610
19.11
34ab
3350a
49a
1.83a
252.65cbd
I/
2/
3/
4/
5/
pLOgZseedZhr. p ic o g .Zseed
7.11b
86d
27
ATP-
.95d
84.47a
37
4Z
76f
2 . 16cd
C.V.%5 /
%
phosZg
Resp.
10
5
5
7
.
Means follo w ed by the same l e t t e r are not s ig n if ic a n t ly d if f e r e n t (P = 0 .0 5 ) according to Neuman-Keuls mean
sep aratio n te s t
Seedling growth ra te
E l e c t r ic a l c o n d u c tiv ity te s t
R e s p ira tio n ra te
C o e ffic ie n t o f v a r i a b i l i t y
21
26
not with speed of emergence (r = 0.74) and first year forage
yield (r = 0.76) .
The high r values are not significant due
■
to few degrees of freedom.
Accelerated aging,
were
in
respiration rate,
and ATP
content
significantly correlated with first year forage
1986
Table
(Table
3.
4).
None
of the
laboratory
yield
tests
were
Simple correlations among standard germination,
accelerated
aging
and
field
performance
variables for four seed lots of Regar meadow
bromegrass - Bozeman, MT. 1985
Laboratory parameters
Field
parameters
Std. germ.
Accel, aging
Speed of emerg. (index)
0.74
0.83
Total emerg. (plants/m)
0.91*
0.97**
Forage yield (kg/ha)
0.76
0.88*
* r (0.05) = 0.88
** r (0.01) = 0.96
correlated with speed of emergence and total emergence.
The
inconsistent relationship among speed of emergence or
total
seedling
years
(Tables
emergence with laboratory tests during both
3
and
4)
suggest that this
association
may
be
effected by the presence or lack of stress conditions in the
field.
First year forage yield was the only parameter which
consistently
reflected
during both years.
vigor differences among
seed
lots
27
Standard germination percentage
with
did not correlate well
first year forage yield (Table 4) during 1986
studies
(r = 0.53) , which agrees with the 1985 data (Table 3). Seed
lots 861 and 862 had standard germination percentages of 96%
and 94% (Table 2 ),
94%
germination,
forage
yield
respectively. However, seed lot 863 with
had
a significantly
than seed lots 861 and 862
higher
(Table
first
year
2).
This
supports previous research by Naylor (1981) and Marshall and
Naylor (1985) who found that Italian ryegrass seed lots with
Table
4.
Simple correlations among standard germination,
vigor tests and field performance variables for
10 seed lots of Regar meadow bromegrass Bozeman, MT. 1986
Laboratory parameters
Std.
germ.
I/
SGR
2/
ECT
3/
AA
Speed
0.38
of emerg.
(index)
-0.09
0.26
0.47
Total
0.37
emerg.
(plants/m)
-0.04
0.14
0.54
0.58
-0.58
4/
Resp
5/
ATP
Field
parameters
Forage
yield
(kg/ha)
*
I/
2/
3/
4/
5/
0.53
r(.05) = 0.60
Seedling growth rate
Electrical conductivity test
Accelerated aging.
Respiration
Adenosine triphosphate
0.71*
0.008
-0.06
0.69*
-0.25
-0.25
0.60*
28
similar
and acceptable standard germination percentages may
not perform similarily in the field.
Seed lots 869 and 8610
had
standard germination percentages of 60% and 49%
2),
respectively.
(Table
Even though these seed lots had the same
number of live seed per meter of row as seed lot 863,
their
first year forage yields were significantly lower (Table 2).
These
data suggest that differences in seed
first
year
forage
viability
does
yield and that
not
improve
vigor
compensating
field
affects
for
seed
of
low
performance
viability seed lots.
Neither
seedling
growth
rate
nor
electrical
conductivity were correlated with first year forage yield in
1986
(Table
4).
Seedling growth rate was different
seed lots (Table 2),
(Table
4)
to
yield.
Seed
but these differences were not related
standard germination or
lot
among
863 and 869 had
first
year
significantly
forage
different
electrical conductivity readings, while the remainder .of the
seed lots did not differ greatly (Table 2). Seed lot 869 was
mechanically
being
stripped
contributed
(1985).
test
damaged, which resulted in the palea and lemma
to higher solute leakage.
found
and
suggested
from the caryopses.
This damage
Marshall and
a poor relationhip between
field
performance
in
may
Italian
the
have
Naylor
conductivity
ryegrass.
that leaching of electrolytes from the lemma
They
and
palea may mask differences among seed lots, or the lemma and
29
palea
might
slow
water
passage
into
caryopses
and
electrolytes out.
First
year forage yield was
significantly
correlated
with accelerated aging (r = 0.71*) and respiration rate (r =
0.69*) (Table 4).
Seed lot 863 had the highest
accelerated
aging value (Table 2) and was significantly higher than seed
lots
861
and
germination
869
and
lowest
seed
862.
These
two
seed
lots
percentages similar to seed lot 863.
8610
performed poorly in the field
accelerated aging values (Table 2).
lots contained high amounts of
could
had
not
be
controlled
with
and
Both
storage
surface
standard
Seed lots
had
the
of
these
fungi,
which
sterilization.
Consequently, seed from these lots deteriorated rapidly when
exposed
the
to accelerated aging conditions.
highest
respiration
rate among
seed
significantly higher than seed lots 861,
(Table
2).
However,
mechanically
injured,
statistically
(1969b)
lot
exhibited
equal to seed lot 863
reported
vigorous,
seed
but
may
Seed lot 863
lots
was
864, 865, and 8610
869,
which
had
respiration
(Table
2).
that mechanically injured seed
have
and
had
increased rather
than
been
rates
Woodstock
was
less
decreased
respiration rates.
There was a significant correlation between ATP content
of
imbibed
seeds and first year forage yield (r
=
0.60*)
(Table 4). Seed lot 863 had the highest ATP content per seed
30
and
was
(Table
significantly different than all other
2).
Seed
lot
869,
which had one of
seed
the
lots
highest
respiration rates, also had a high ATP value (Table 2). High
respiration rates may contribute to the high ATP
of
seed
lot
correlation
869,
between
(Appendix-Table
correlations
and
for
impractical
the
significant
respiration rate and ATP (r
7).
Ching
between
seedling
techniques
which is supported by
production
size
(1973)
observed
0.71*)
significant
ATP content of imbibed ryegrass
and
measuring
weight.
ATP
Even
content
as a seed vigor test,
though
in
seed
present
seeds
may
be
its requirement for seed
germination and early seedling growth makes it an
biochemical
=
important
tool for evaluating seed vigor and providing an
explanation for the phenomenon of seed vigor.
A multiple stepwise regression model building procedure
was
used
associated
was
to
determine which 1986 seed
with first year forage yield.
selected
vigor
tests
Accelerated aging
first and was most closely related
year
forage yield (R = 0.71*)
also
had
(Table 5).
the highest correlation with
were
to
first
Accelerated aging
first
year
forage
yield during 1985 preliminary studies (Table 4). Respiration
rate
strengthened
the above relationship and was the
other variable added to the model (R = 0.78**)
only
(Table 5).
These data suggest that one vigor test is not enough to
measure overall seed quality.
A combination of tests, which
31
measure
both physiological and biochemical aspects of
seed
vigor should be utilized. For Regar meadow bromegrass, first
Table
5.
'
Summary of multiple stepwise regression comparing
first year forage yield with vigor tests for 10
seed
lots
of
Regar meadow
bromegrass Bozeman, MT. 1986
2
Step
Variable
1
Accelerated
aging (A.A.)
0.51
2
Respiration
(Resp.)
0.61
year
and
R
Regression equation
y = 3355 + 22(A.A.)
y = 2.794 + 15 (A.A.) + 1332 (Resp.)
forage yield was closely related to accelerated
respiration
imbibed
vigor.
seeds
rate.
may
Adenosine
triphospate
content
be a useful biochemical index
x
aging
of
of
seed
32
LITERATURE CITED
i'
33
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APPENDIX
41
APPENDIX
Table
6.
Simple correlations among standard germination,
accelerated aging and field performance variables
for four seed lots of Regar meadow bromegrass Bozeman, MT. 1985
Field parameters
Speed
of emerg.
I
Total
emerg.
2
Laboratory parameters
Forage
yield
3
Std.
germ.
4
I
1.00
2
0.94*
1.00
3
0.96**
0.94*
1.00
4
0.73
0.91*
0.75
1.00
5
0.83
0.96**
0.88*
0.97**
* r (0.05) = 0.88
** r (0.01) = 0.96
Accel.
aging
5
1.00
Table 7.
Simple correlations among standard germination,' vigor tests, and field performance variables
for ten seed lots of Regar meadow bromegrass - Bozeman, MT, 1986.
Speed
o f emerg.
I
F ie ld parameters
T o ta l
emerg.
2
Forage
y ie ld
3
S td .
germ.
4
SGR
5
Laboratory parameters
A ccel.
2/
ECT '
aging
6
7
3/
R e s p .7
8
I
1.00
2
0 .9 3 * *
1.00
3
0 .13
0 .12
1.00
4
0 .3 8
0 .3 8
0.53
1.00
5
- 0 .0 8
- 0 .0 4
0 .58
0 .43
1.00
6
0.26
0 .1 3
- 0 .5 8
- 0 .2 7
- 0 .5 0
1.00
7.
0 .47
0 .5 3
0 .7 1 *
0 .7 2 *
0 .52
-.0.54
1.00
8
0.008
-0 .0 6
' 0 .6 9 *
0.51
0 .6 1 *
-0 .3 5
0 .6 1 *
1.00
- 0 .2 5
0 .6 0 *
0 .20
0 .42
- 0 .6 6 *
0 .34
0 .7 1 *
9
*
**
I/
2/
3/
- 0 .2 4
r ( 0 . 0 5 ) = 0 .60
r ( 0 . 0 1 ) = 0 .73
seedlin g growth ra te
e le c t r ic a l c o n d u c tiv ity te s t
r e s p ira tio n ra te
ATP
9
1.00