Published May, 1990
Seed Production Environment Influence on the Base Temperature
for Growth of Sorghum Genotypes
D. J. Lawlor,* E. T. Kanemasu, W. C. Albrecht, III, and D. E. Johnson
ABSTRACT
The base temperature for growth is an important element in plant
growth models that utilize heat accumulation methods. The effect of
location (environment) on the base temperature for growth of sor-
ghum (Sorghum bicolor (L.) Moench) genotypes was studied. Seeds
of six sorghum genotypes were obtained from three locations: Man-
hattan, KS, College Station and Lubbock, TX. The seeds were
placed on moist germination blotter paper on a one-way thermal
gradient plate. Observations were made at least daily for 3 wk to
determine the number of days required for (i) germination (i.e., pro-
trusion of the radicle from the seed), (ii) root elongation to 10 mm
or longer, and (iii) shoot elongation to 10 mm or longer. The base
temperature for germination of each genotype was then determined
as the x-intercept from regressing the rate of germination (i.e., the
reciprocal of the number of days required to germinate) vs. the mean
temperature. A similar regression was used to determine the base
temperatures for shoot elongation and for root elongation of each
genotype. The values of base temperature for shoot and root elon-
gation were similar for the genotypes studied, and were consistent
with the values of base temperature for emergence of sorghum re-
ported in the literature; whereas the base temperature for germi-
nation values were up to 4.6 oc lower than reported values. Results
indicated that: the base temperature for shoot elongation is not af-
fected, the base temperature for root elongation is somewhat af-
fected, and the base temperature for germination is more affected
by the location (environment) of seed production.
W.C. Albrecht, III, and D.E. Johnson, Evapotranspiration Lab.
Dep. Agronomy, Kansas State Univ., Manhattan, KS 66506; D.i
Lawlor, Texas A&M A¢c. Res. and Extension Center, Route 2
Box 589, Corpus Christl, TX 78410; and E.T. Kanemasu, Dep. of
Agron., Univ. of Georgia, Griffin, GA 30223-1797. Contribution
no. 88-134-J, Kansas Agric. Exp. Stn., Manhattan, KS 66506. Re-
ceived 4 May 1988. *Corresponding author.
Published in Agron. J. 82:643-647 (1990).
644 AGRONOMY JOURNAL, VOL. 82, MAY-JUNE 1990
T HE BASE TEMPERATURE for growth is an important
element in plant growth models that utilize heat
accumulation methods. Errors in the base temperature
used in a model may lead to erroneous estimations of
phenological events and yield, particularly in cool
areas. From a production standpoint, it is important
to have plants that will grow at low temperatures in
areas with high altitude, a short growing season, or a
cool spring. . (McBee et al., 1983).
Several studies have found that cultivars of a given
crop have the same base temperature (Tb) for growth,
for example, spring wheat (Triticum aestivum L.) (del
Pozo et al., 1987), pearl millet (Pennisetum typhoides
S. & H.) (Garcia-Huidobro et al., 1982), and sorghum
[Sorghum bicolor (L.) Moench] (Kanemasu et al.,
1975). In contrast, Thomas and Miller (1979) found
significant differences in base temperatures for ger-
mination among sorghum cultivars, with base tem-
peratures ranging from 6.9 to 13.4 oc.
Garcia-Huidobro et al. (1982) found that two cul-
tivars of millet (a modern hybrid and an indigenous
variety of unknown source) showed a similar response
to temperature. Values of Tb and Tm (maximum tem-
perature for growth) were the same for both, and T0
(optimum temperature for growth) differed by only
about 2 oc. The authors suggested that all seeds (of a
species) have a common base temperature but need
different periods of thermal time to germinate. Using
six hybrid millet cultivars, Ong (1983) found a similar
base temperature (11.3-13.2 °C) for all the vegetative
processes examined (i.e., seedling emergence, leaf ini-
tiation, leaf appearance, and tillering from sowing).
Little work, however, has been done on the possible
effect oflocation (environment) of seed maturation on
the base temperature for growth of crop genotypes.
Burris ( 1977) found that location of production and
female parentage had a significant effect on germina-
tion and early shoot and root dry weight of corn (Zea
mays L.). However, an early frost occurred during his
experiment, which may have affected his results.
Mohamed et al. (1985) studied the effect of tem-
perature during seed development on germination
characteristics of pearl millet seeds. Their results
showed that cardinal temperatures (Tb, T 0 , and T m)
were unaffected by the environmental temperature
during seed development (i.e., 22-31 °C) and growth
(i.e., grain-fill), whereas these conditions did affect
seed size and, subsequently, germination rate and seed
viability. However, base temperatures for germination
were found to be related to adaptation within sorghum
species (Thomas and Miller, 1979), with the base tem-
perature of temperate hybrids (temperate types) being
about 10 oc and those of tropically adapted hybrids
(tropical types) about 7 oc. Mann et al. (1985) found
that sorghum cultivars which are adapted to temper-
ature areas showed higher G50 values (G 50 = the low-
est temperature at which 50% of a specific seed lot will
germinate in a specified time period; i.e., the base tem-
perature for germination) than the cultivars adapted
to more tropical environments. They also found that
a lower G 50 value corresponded with a lower Tb. From
an ecologist's point of view, the ability of temperate
or more northern types to germinate at a higher tem-
perature than those with tropical or more southern
types might be indicative of an adaptation property
to avoid germination until the environmental condi-
tions are warm enough to ensure growth to maturity
(Abdalla and Miller, 1982). Other data (Mann and
Miller, 1984) suggest that Tb of the temperate and
tropical sorghums varies with location (environment)
of seed maturation and might be related to the sugar
content of the seed, because tropical types tend to
have higher stem sugar contents than temperate types
The objectives of this study were to determine the
base temperature for growth of several genotypes of
sorghum produced at each of three locations and to
determine whether the base temperature for growth
varies among genotypes and/or locations (environ-
ments).
MATERIAlS AND METHODS
Seeds of six genotypes of sorghum were obtained that had
each been field produced in 1985 at three locations: Man-
hattan, KS, and College Station and Lubbock, TX. The six
genotypes were: BT X 378; BT X 398; BT X 399; BT X
623; RT X 7000; and T X 09. The genotypes were of the
same relative maturity at the three locations. (The seeds were
provided by Dr. J.A. Mann, Texas A&M University, College
Station; Dr. D.T. Rosenow, Texas Agricultural Experiment
Station, Lubbock; and Dr. P.J. Bramel-Cox, Kansas State
University, Manhattan).
The seeds were placed, embryo side up, on moist seed
germination blotter paper on a one-way thermal gradient
plate. Distilled water was applied to the blotter paper, as
needed to prevent dehydration during the duration of the
experiment. Room temperature (20 oq distilled water was
applied to the warm end, and refrigerated distilled water was
applied to the cold end. The thermal gradient plate consisted
of an aluminum plate (0.91 m wide by 1.22 m long by 25.4
mm thick), which was cooled at one end and heated at the
other end to establish and maintain a uniform temperature
gradient by pumping cold/hot water-ethylene glycol mixture
through 6.4-mm-copper tubing attached to the underside of
the aluminum plate at either end. The temperature range
studied was 0 to 30 °C. Once established, the thermal gra-
dient was linear from cold end to warm end and was held
constant for the duration of the experiment. A plexiglass
cover (1.08 m long by 0.91 m wide by 6.4 mm thick)
mounted 13 mm over the plate aided in water retention. A
25 mm layer of foam plastic insulation enclosed the thermal
gradient plate/plexiglass on all sides. A 13-mm-thick ply-
wood box enclosed the insulation foam. The box was tilted
at a slope of approximately 3o so that water would run from
the cold end to the warm end to help maintain the temper-
ature at the cold end. The entire system operated in a con-
stant temperature room to minimize ambient air influences
on the water bath temperatures. Copper-constantan ther-
mocouples were used to monitor the temperature of the ther-
mal gradient plate at 25.4-mm intervals from cold end to
warm end as well as at several locations toward the outer
edges of the plate. The data were recorded on cassette tape
and on paper printout (CR5, Campbell Scientific Inc., Logan,
UT) 1 at hourly intervals. There was generally less than a
0.5 oc variation in temperature across the plate.
The experimental design was completely randomized with
two replications on the plate at the same time. Seeds of a
given genotype were placed at 25-mm intervals in a line from
the cold end to the warm end (42 seeds, 1 per 25 mm = 1
replication); rows of the various genotypes were placed at
25-mm intervals across the plate. Observations were made
at least daily for 3 wk to determine the number of days
1 Company names and trade names are included for the benefit
of the reader and do not imply any endorsement.
 LAWLOR ET AL.: SEED PRODUCTION ENVIRONMENT INFLUENCE ON SORGHUM 645

0.6 X X
Table 1. Adaptation type, and thermal units (TU) and mean base
temperature (T.s, T.,R, T.G) required for shoot elongation, root
0.5 elongation, and germination of six sorghum genotypes produced
X
z at three locations.
0
U.i= - 0.4 Shoot
elongation
Root
elongation Germination
0<('7 Adaptation
~~ ~ 0.3 Genotype typet r.s TV T.R TV T.G TV
<C ~ 0
a:wa:- 0.2 oc oc d-1 oc oc d-1 oc oc d-1
BT X 623 TA 10.6 75.2 9.7 44.2 5.3 31.9
<!J T X 09 TA 10.3 68.5 10.5 39.5 5.9 27.8
0.1
BT X 399 TE 10.0 92.7 11.0 51.0 9.1 27.1
RT X 7000 TE t t 9.4 55.6 7.4 30.2
0'~--~-+------+------+------+-----~ BT X 378 TE 9.9 100.5 9.3 70.4 8.7 38.3
4 8 12 16 20 24 BT X 398 TE 9.7 90.1 8.6 65.2 8.5 31.9
0
MEAN TEMPERATURE ( C) LSD (0.05) 1.3 15.5 1.8 15.2 2.3 9.4
(NS) (NS)
Fig. I. Rate of germination of sorghum vs. mean temperature of the
seeds for genotype BT X 623. t Adaptation type as classified by Texas Agric. Exp. Stn. sorghum breeders:
TA = tropically adapted; TE = temperately adapted.
t Insufficient data to determine T.S and TV.
required for (i) germination (i.e., protrusion of the radicle
from the seed), (ii) root elongation to 10 mm or longer, and Table 2. Mean base temperature (Tb) and number of TU required
(iii) shoot elongation to 10 mm or longer. Seeds were dis- for observational periods of growth of five sorghum genotypes pro-
carded after the shoot had elongated to 10 mm or longer. duced at three locations.
The relation between number of days (duration) to reach
a given observational stage [e.g., (i), (ii), or (iii) above] and Shoot Root
elongation elongation Germination
mean temperature is curvilinear. However, the rate of de-
velopment, defined as the inverse of the time to reach a Location Latitude r.s TV r.R TV r.G TV
certain developmental stage, increases linearly within a de- oc ocd-1 oc ocd-1 oc ocd-1
fined range of temperatures (Angus et al., 1981; Garcia-Hui-
dobro et al., 1982; Kanemasu et al., 1975; del Pozo et al., College Stn., TX 30°36' 10.7 74.8 10.7 48.6 8.3 28.3
Lubbock, TX 33°36' 9.7 93.1 9.9 53.1 8.2 30.4
1987). Data for most species suggest a linear relationship Manhattan, KS 39°09' 9.9 88.3 8.7 61.4 5.9 34.9
(Angus et al., 1981). When the rate is linear, extrapolation LSD (0.05) 1.0 12.0 1.3 10.7 1.6 NS
defines a base temperature, Tb, at which the rate is zero so Overall mean 10.1 85.4 9.8 54.3 7.5 31.2
that the thermal response of germination (or root or shoot
elongation) can be expressed as
1/t = (T - Tb) b The base temperature differences between the two
stages of growth (root elongation and shoot elongation)
where t is the time required to reach a particular develop- were small (0.2-1.1 °C}. The TbS and TbR values ob-
mental (or observational) stage, T ( C) the mean tempera-
0
tained were consistent with the range of base temper-
ture during this period, Tb the base temperature, and b the atures for sorghum found in the literature (e.g., 7-
reciprocal of the thermal time required for the process to
occur (Monteith, 1977). 13 oc, Angus et al., 1981; Mann et al., 1985; Thomas
The base temperature for germination ( TbG) of each geno- and Miller, 1979).
type was then determined as the x-intercept from regressing The genotype rankings for TbG were different from
the rate of germination (i.e, the reciprocal of the number of those of TbS and TbR (Table 1). The TbG values also
days required to germinate) vs. the mean temperature (of were up to 4.6 oc lower than values for TbR. This
the plate where the seed was located) (Fig. 1). The number agrees with other studies using thermogradient plates,
of thermal units (TU) required for germination was then which have found that TbG is lower (up to 3 oq than
obtained by taking the inverse of the slope of the regression laboratory or field determination of Tb for emergence
line. A similar regression was used to determine the base of numerous field crops (Angus et al., 1981), vegetable
temperature for shoot elongation (TbS) and for root elon-
gation (TbR) of each genotype. species (Bierhuizen and Wagenvoort, 1974), tomatoes
Data were analyzed by using analysis of variance (AN- (Lycopersicum esculentum) (Thompson, 1974), and
OVA) techniques (SAS Institute, Inc., 1982). Differences re- sorghum (Kanemasu et al., 1975).
ported are those that were significant at P = 0.05 based on Angus et al. (1981) pointed out that not all the seeds
the least significant difference (LSD) test. which germinate at a very low temperature develop
into emerged seedlings because of losses due to path-
RESULTS AND DISCUSSION ogens and exhaustion of seed reserves. This accounts
for the low values found fQr TbG, compared to TbS
Genotypic Differences
and TbR. The terminology used by various authors
There were no significant differences in TbS (mean also may be misleading. For example, Garcia-Hui-
TbS = 10.1 oq among genotypes, averaged across lo:.. dobro et aL (1982) defined a seed as germinated when
cations (Table 1). Genotype RT X 7000 from Man- the radicle was 10 mm (or longer), thus their definition
hattan, KS, produced insufficient data to determine of TbG is not germination as we have defined it, but,
TbS, so only five genotypes are discussed for TbS. The developmentally corresponds to our definition of TbR.
number ofTU required for shoot elongation were sig- This supports our use of T~ (and TbS) over TbG.
nificantly different among genotypes. The ranking of There were no significant differences among genotypes
the genotypes for TbR (Table 1) was similar to that for TU required for germination.
for TbS, although T~ showed significant differences The tropical genotypes (BT X 623 and T X 09) had
among genotypes (as TU required for root elongation). TbG values significantly lower than three of the four
646 AGRONOMY JOURNAL, VOL. 82, MAY-JUNE 1990

Table 3. Mean base temperature ( 0 C} for germination (T.G), root elongation (T.,R) and shoot elongation (Tt$) of six sorghum genotypes
produced at three locations (CS = College Station, TX; Lub = Lubbock, TX; and Man = Manhattan, KS).
r.s T.R T.G
Genotype cs Lub Man cs Lub Man cs Lub Man
T X 09 9.6 10.9 10.6 10.8 11.8 9.0 7.4 3.5 7.0
BT X 378 12.1 9.4 8.4 12.2 8.9 6.8 9.4 11.4 5.4
BT X 398 10.4 9.2 9.5 8.2 9.8 7.7 9.7 10.1 5.7
BT X 399 9.9 9.4 10.8 11.3 10.3 11.5 8.6 8.9 9.9
BT X 623 11.7 9.9 10.3 11.8 7.9 9.4 6.2 6.2 3.5
RT X 7000 t t t 10.0 10.7 7.6 8.9 9.3 3.9
LSD (0.05) 2.2 3.1 4.0
Mean 10.7 9.7 9.9 10.7 9.9 8.7 8.3 8.2 5.9
t Insufficient data to determine T.S-
temperate genotypes (Table 1), which supports the (nonsignificant) TbG over locations. Genotypes BT X
findings of Mann et al. (1985) that tropical types have 378 and BT X 398 from Manhattan had a significantly
lower TbG's than temperate types. However, tropical lower TbG than BT X 398 from Lubbock (although
genotypes had higher TbR values than three of the four not a significantly different value than College Sta-
temperate genotypes and the highest TbS values tion), whereas R T X 7000 from Manhattan had a sig-
(though not significantly higher than any of the tem- nificantly lower TbG than R T X 7000 from College
perate genotypes). This indicates that the base tem- Station or Lubbock.
perature for germination (as we have defined it) may
not be a true indication of the base temperature for CONCLUSIONS
growth of a species, although it may still be of use to Values of TbS and TbR were similar for the geno-
distinguish between tropical and temperate genotypes. types studied, ranging from 8.6 to 11.0 °C, and were
consistent with the values of base temperature for
Location Differences emergence of sorghum reported in the literature, while
For each location, TbS and TbR were similar (dif- TbG values were as much as 4.6 oc lower than re-
fering by 0.0-1.2 °C), whereas TbG was low (1.5- ported values. For use in plant growth models, a base
2.8 oc lower than TbS or TbR; Table 2). Each of the temperature calculated from a more advanced stage
three traits showed significant differences in Tb among of growth than germination is suggested because, as
locations; however, TbS was significantly different Angus et al. ( 1981) noted, not all seeds which germi-
among locations only because one of the five genotypes nate at very low temperatures develop into emerged
(i.e., BT X 378) was highly significantly different seedlings because of losses due to pathogens and ex-
among locations while the other four were not signif- haustion of seed reserves. The use of TbS or TbR over
icantly different among locations or among genotypes TbG values would thus lead to less errors in estima-
(Table 3). Thus the difference in TbS among locations tions of phenological events and yield.
in Table 2 is due to a single hybrid. Genotypes from The TbS is not affected, TbR is somewhat affected,
College Station had the highest mean Tb at each stage and TbG is more affected by the location (environ-
of growth, whereas those from Manhattan had the low- ment) of seed production. While tropical genotypes
est (though not significantly lower than those from had significantly lower TbG values than temperate
Lubbock for TbS or TbR). genotypes, TbR and TbS values were not significantly
The number ofTU required for shoot and root elon- different between tropical and temperate genotypes,
gation was significantly higher for Manhattan geno- indicating that separation of these genotypes by base
types than for those from College Station, whereas temperatures may not be plausible at later stages of
Manhattan genotypes did not differ significantly from growth.
Lubbock genotypes (Table 2). The TU required to
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