Published January, 1968

Effect of Seed Pod and Temperature on Speed of Germination and Seedling

Elongation of Sainfoin ( Onobrychis vici~folia Scop.)t
2A. E. Carleton, C. S. Cooper, and L. E. Wiesner

ABSTRACT concluded that a water-soluble inhibitor was not pres-

Optimum temperatures for germination of sainfoin ent in the pod. In contrast, Cavassa (3), cited by Op-
seed and for seedling growth were determined in con- penheimer (6), found that the sainfoin pod hindered
junction with studies of the effect of the presence or imbibition of water and contained inhibiting sub-
absence of the seed pod. Optimum temperature range for
germination of sainfoin was from f5 to 20 C. A higher stances which delayed emergence of the radicle and
range of optimum temperature, 20 to 30 C, was found hampered its development.
for seedling growth. The presence of pods at all tempera-
tures reduced speed of germination and speed of seedling
elongation. Pods on the seed slowed water absorption PROCEDURE, RESULTS, AND DISCUSSION
by 4 to 5 hr. A water soluble inhibitor which slowed the
speed of germination of shelled seed was detected. This Effect of Pod on Germination and
inhibitor was readily removed from the pod by washing. Seedling Elongation
Mechanical restriction of germination by the pod ap-
peared to be a minor factor in the total effect of the pod Replicated lots of shelled and unshelled sainfoin seed were
on speed of germination. There was no difference in germinated in the dark over a temperature range of 15 to 35 C.
field emergence rates of sainfoin seedlings from shelled Seeds were placed in 4 X 4-inch plastic trays on blotter paper
and unshelled seed. moistened with 30 ml of dlstilled water. The percentage germ-
ination of seeds and total seedling length (cotyledons to tip of
Additional index words: effects of moisture and in- radicle) were determined daily over an 8-day period. Speed of
hibitors., germination (SG) was calculated by the method of MaGuire (5)
where:
~-
T HEScop.)
legume sainfoin
is attracting
(Onobrychis viciaefoIia
widespread interest in the
SG ~= (no.
~y~ ~- ~r~-~
normaI seedlings.... -~ ~-a~no. tonormal
final seedlings’~
count

Northern Rocky Mountain region. In preliminary Speed of elongation was calculated by the formula:
testing the variety ’Eski’ appears to be adapted as a
range legume to areas with 13 or more inches of SE = mean seedling length -b ....
~-~ to first count
precipitation. Hay yields of sainfoin were similar to
alfalfa in short-term irrigated hay trials in two-cut mean increase in length from previous count’~
areas (1). Sainfoin yielded more than alfalfa in
area where short water supply limited production to
days to last count )
The effect of temperature and of the pod on speed
one cutting (1). Sainfoin is not knownto cause bloat.’ of germination is shown in Fig. 1. Germination of
Potential for bloat is similar to birdsfoot trefoil (Lotus shelled seed was most rapid at 20 C. The next most
corniculatus L.) based on in vitro foam formation of rapid germination temperature was 15 C. Rapidity
extracts from each species (2). of germination of shelled seed decreased as tempera-
Since sainfoin has not been investigated extensive-
ly in the United States, information is needed concern-
ing its culture. The present study is concerned with the 55-
effect of the seed pod and temperature upon germina-
tion and seedling elongation. Sainfoin seed matures in
one-seeded pods, described by Piper (7) as "brown, 5O
indehiscent, lenticular, and reticulated on the sur-
face." The pod is bean-shaped and bilaterally com-
pressed, and it has a rough net-veined appearance. The 25
~eed itself is kidney-shaped, varying from yellowish
green to dark brown in color. The seed is much
larger than seed of our commonforage legumes. A ~o
pound of seed in the pod contains about 18,800 seeds
and a pound of shelled seed about 26,500 seeds (4).
Thus, the pod comprises 29% of the seed weight.
The effect of the pod upon germination was pre-
viously studied by Thomson (8). He concluded that I0,
lower germination was due to the failure of the radi-
cle from weaker seed to penetrate the pod. He also .5-
z Cooperative investigations of the Montana Agricultural Ex-
periment Station and the Crops Research Division, Agricultural
Research Service, USDA.Published with the approval of the
¯ Director of the Montana Agr. Exp. Sta. as Journal Series 814.
Received June 6, 1967. GERMINATION TEMPERATURE (,C)
~Assistant Professor; Agronomist, Crops Res. Div., ARS, USDA;
and Seed Analyst, Montana State Univ., Bozeman, Mont., re- Fig. l. Effect of germination temperature and seed condition
spectively. on speed of germination.

81
82 AGRONOMYJOURNAL, VOL. 60, JANUARY-FEBRUARY 1968

tures increased above 20 C. Germination of seed in
the pod was similar at a range of 15 to 23 C and de-
creased at 30 and 35 C. At all temperatures, seed
in the pod was slower to germinate than shelled seed.
At the end of 8 days total germination of both shelled
and unshelled seed was less at 30 and 35 C (Fig. 2).
Total germination of shelled seed was greater than un-
shelled at all temperatures. This difference may have
been due to a difference in seed quality. Immature
seeds were readily detected and removed from shelled
seed but could not be detected in seed in the pod.
Temperature optimums for seedling growth were
.different from those for germination (Fig. 3). Optimnm
temperatures for speed of seedling elongation ranged
from 20 to 30 C for shelled and from 15 to 30 C for
seed in the pod. At all temperatures, speed of seedling
elongation of Shelled seed was more rapid. Conse-
quently, total seedling length at 8 days was general-
ly greater for seedlings from shelled seed (Fig. 4).
The decrease in speed of germination and of seed-
ling elongation caused by the pod (Fig. 1 and 3)
could be due to several factors; namely, (1) a slowed
rate of water absorbtion, (2) the presence of an in-
hibitor, and (3) mechanical restriction. These factors
were studied in .subsequent experiments.
Effect of Pod on Rate of Water Absorption
Replicated 100-seed lots of shelled and unshelled sainfoin
seed were germinated at 20 C. The germination procedure was
the same as in the previous study. At 20-minute intervals for
the first hour, and 1-hour intervals the next 15 hr, replicates
of each seed type were relnoved. Pods were re~noved from un-
shelled seed and the amount of moisture absorbed by each seed
type was determined.
Water absorption by shelled seed was very rapid
(Fig. 5). After 4 to 5 hr the water content rose
between 50 and 55%, where it remained for the dura-
tion. of the test. In contrast, water content of seed in
the pod did not approach that of shelled seed until 9
to 10 hr. The differences of 4 to 5 hr in the time
needed to reach maximumabsorption is not likely to
be of much importance, given adequate water for
germination. However, with summer showers and
rapid drying, this lag might be enough to prevent seed
from germinating at a time when seedbed moisture
is not adequate to sustain the seedlings.
IOO-
90-

80’
IOO-
HEL
~ SHEL LED g ~’o-
90- "i-
I--
o 60-

o 50-
~ 70- Z
tu _J
~ 40-
~ 6o-
IN THE POD
~0-
50-

20-
40-
15 20 23 30 35
15 20 23 30 35 GERMINATION TEMPERATURE(C)
GERMINATION TEMPERATURE(C)
Fig. 4. Effect of germination temperature and seed condition
Fig. 2. Effect of germination temperature and seed condition upon seedling length after 8 days.
on percent germination after 8 days.

z 25

z ~ 40 /N THE
o
.J ~-35-
"~ 15

r~ I0
o 20-~
uJ

,, 5
0 l=.+si=_+ 1.8 IO-

uJ 0 I I I I
Q. 0 I I I I I I I I I I I I I I I
u) 15 20 23 30 35
GEI~IMINATION TEMPERATURE (C) H~ IN ~EOMIN&TO~
Fig. 3. Effect of germination temperature and seed condition
on speed of seedling elongation. ¯ ig. ~. Water uptake of shelled and ~sheIled sa~fo~ seed.
 CARLETON ET AL.: GERMINATION OF SAINFOIN 83

The Presence of an Inhibitor I00- ------- PODSREMOVED

Pods from 100 sainfoin seeds were soaked for 18 hr in 25 ml 90- PODSOPEN, WASHED
of distilled water. The water was’ then strained from these seeds --.-- PODS OPEN, NOT WASHEDI
and was used to moisten blotter paper in 4 X 4-inch plastic
trays upon which shelled sainfoin seed was germinated at 20 C. ozOO--TO .-- PODS /NTACT..." //~/f
Shelled and unshelled seed, moistened with 25 ml of water, were
included as controls.
The number of seeds
speed of germination
Each treatment
germinated
calculated
was replicated
each day was counted
as previously
three times.

described.
and
~ ~o
~ 50 ,,.///
The addition of the water extract from the sain-
~ ~o. .
loin pods slowed rate of germination. Speed of germi-
nation (SG) of shelled seed, shelled seed with extract
added, and seed in the pod was 70.1, 50.1 and 28.9,
respectively. These values were significfintly dif- , , , ,
ferent. The addition of the extract did not slow 12 24 36 48 60 72 84 96 108 120 132
germination to the rate observed for seed in the pod. HR IN GERMINATOR
This was probably due to a dilution effect from spread- ¯ i~. 6. ]nflucn~c o[ ~our sec~ trea~ents ~pon germina~o~ ~ate
ing the extract over the total blotter surface in the o~ sMn~oin.
germinating dish.
In a second study, a sample of unshelled sainfoin seed was
soaked in water for 18 hr and then air-dried for 7 days. The I00-
sample was then divided and one-half was soaked a second time
for 18 hr and dried for 7 days. Replicated seed lots from these
two samples were germinated in comparison with unsoaked 90-
shelled and unsoaked, unshelled seed.
Soaking seed did not reduce total germination 80-
(Table 1). Seed in the pod soaked twice germinated
more rapidly than unsoaked unshelled seed (Table 70-
1). Germination appeared to be faster for seed soaked
twice than for seed soaked once although the differ- ’1- 60-
ence was not significant. Seed in the pod soaked twice I--
germinated as rapidly as shelled seed. These data sug- Z
gest the presence of a water-soluble germination in- 50-
hibitor in the sainfoin seed pod.
A third study was conducted in which sainfoin seed was sub- 40-
jected to the following four treatments: (1) pods removed; (2)
pods opened but left on seed; (3) pods opened, removed from
seed, washed for 16 hr, oven-dried, and replaced on seed; and 30-
(4) seeds left undisturbed in the pods. Eight replications of
100 seeds each were used for each treatment. After treatment, 20-
seed lots were placed in 4 X 4-inch plastic trays on blotter
paper, moistened with 30 ml distilled water and germinated at
20 C. The percentage germination was determined at 12-hr in- I0-
tervals over a 132-hour period.

Removal of pods resulted in the earliest and most

rapid germination (Fig. 6). Whenpods were removed, 3 4 5 6 7 8
washed, and replaced on the seed, germination rate TIME IN GERMINATOR (days)
was only slightly less than when pods were perma- Fig. 7. Effect of the presence of the seed pod on seedling elonga-
nently removed. Most of the difference between ger- tion at 20 C.
mination rate of this treatment and of shelled seed
can be accounted for by difference in water-absorp-
tion rate due to the presence of the pod (Fig. 5). Open- These data also suggest that a water-soluble inhibitor
ing the pod, but leaving it in place on the seed, did is present in the pods, causing slower seed germina-
not affect germination differently from leaving the tion and seedling growth.
seed undisturbed in the pod. Thus, mechanical im-
pedence of germination by the pod was not a factor Effect of Pods on Field Emergence
in germination of these seed lots. Both the unshelled
seed and seed with the pods opened but left on ger- Speed of seedling emergence from shelled and un-
minated more slowly than the shelled seed or seed shelled seed was measured over a wide range of seed-
which had had the pod removed, washed, and replaced. ing rates at three Montana locations. Speed of emerg-
ence was calculated by substituting emergence num-
bers for germination numbers in the formula of Ma-
Table 1. Effect of soaking on total germination and speed of Guire (5). The mean values for 24 replications
germination of sainfoin seed. shelled and unshelled seed were 8.98 and 8.07 at Boze-
Speed of germtnatlon~
Seed condition and treatment Total % germination* man, 11.06 and 10.78 at Huntley, and 8.08 and 7.29 at
In pod (control) 65.7 a* 24.3 a*
In pod (soaked once) 74.3 ab 34. 3 ab Moccasin. Differences were not observed in speed of
In pod (soaked ~wice)
Shelled (control)
79. Obc
88.5 c
43. lb
44.4b emergence or in stands obtained from the two types of
Values not followed by letters in commonare significantly different at the . 05 level seed. The absence of differences under field conditions
of probability, t Calculated by method of MaGulre(5). could be due to the rapid dissipation of the water-
84 AGRONOMY JOURNAL, VOL. 60, JANUARY-FEBRUARY 1968

soluble inhibitor as a result of frequent rain showers

at all locations. The presence of an inhibitor, how-
ever, could retard germination under field conditions,
if adequate moisture were not present to rapidly re-
move it. In addition, the retarding effect of the in-
hibitor on seedling elongation (Fig. 7) could be de-
trimental to seedling survival under conditions of rap-
id soil drying. Under these conditions the shallower
root penetration of seedlings from unshelled seed
might result in their failure to reach moisture avail-
able at greater soil depths.
From our studies we conclude that the sainfoin seed
pod slows water absorption, and we believe that it
contains a water-soluble inhibitor which slows both
speed of germination and speed of seedling elongation.
Under our experimental conditions, however, these
effects were of little significance in field establish-
ment of seedlings.