Allen Press

Weed Science Society of America

Wild Carrot Seeds: Germination and Dormancy

Author(s): H. M. Dale and P. J. Harrison
Source: Weeds, Vol. 14, No. 3 (Jul., 1966), pp. 201-204
Published by: Weed Science Society of America and Allen Press
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 DALE AND HARRISON:
responded to diazotization and coupling with a pink
color. Thus it appeared that soybean roots reduced
aromatic nitro groups to aromatic amino groups.
ACKNOWLEDGEMENTS
This research was supported in part by the supply of
amiben and derivatives as well as a grant from Amchem
Products, Incorporated, Ambler, Pennsylvania.
LITERATURE CITED
1. ANDREAE, W. A. and N. E. GOOD. 1957. Studies on 3-indole-
acetic acid metabolism. IV. Conjugation with aspartic acid
and ammonia as processes in the metabolism of carboxylic
acids. Plant Physiol. 32:556-572.
2. ASHTON, F. M. 1958. Adsorption and translocation of radio-
active 2,4-D in sugar cane and bean plants. Weeds 6:257-262.
3. BAKER, R. S. and G. F. WARREN. 1962. Selective herbicidal
action of amiben on cucumber and squash. Weeds 10:219-
224.
4. COLBY, S. R., G. F. WARREN, and R. S. BAKER. 1964. Fate of
amiben in tomato plants. J. Agr. and Food Chem. 12:320-321.
5. . 1965. Herbicide metabolism: N-glycoside of ami-
ben isolated from soybean plants. Science 150:619-620.
Wild Carrot Seeds: Germination and Dormancy'
H. M. DALE and P. J.
Abstract. Germination of wild carrot (Daucus carota, L.) was not
confined to any season but was correlated with abundant soil mois-
ture in field studies. In laboratory studies, germination per-
centage for any seed lot increased with time as dormancy was over-
come. Further, the germination was not clearly correlated with seed
weight, light, or pH, but temperature and the availability of
water were clearly significant. Dormancy was due to the endosperm
which surrounds the embryo. When part of the endosperm covering
the tip of the radicle was digested or broken off, germination was
increased nearly to the potential for a given lot of seed.
INTRODUCTION
BSERVATIONS made throughout the growing season in
neglected fields and pastures revealed that the ap-
pearance of seedlings of wild carrot (Daucus carota L.) is
not confined to one part of the season. As many as four
flushes of germination have been observed. Harper and
Sagar (5) have shown that seeds of field buttercups
(Ranzinculus acris L.) sown in the autumn germinate in
two groups-one in the autumn and one in the spring.
Such a variation within a seed sample confers on butter-
cup an insurance against destruction of the whole popula-
tion in an adverse season.
Refinements in chemical weed control techniques de-
pend not only on the selectivity of the chemical used but
also on the timing of the application which must be made
when the plant is most susceptible. With delayed dor-
mancy and irregular germination, a weed population may
'Receive(d for ptublicationi Auiguist 30, 1965. Contribution from
I)epartment of Botany, Onitario Agricultural College, IUniversity of
(Guelph, (Guielph, Ontario, Canada.
2Associate Professor and Graduate Student. The latter's present
address: Mfantsiman School, Box 14, Saltpond, Ghana, West Africa.
201
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WILD CARROT SEEDS
6. . 1966. Fate of the amide and methyl ester of amiben
in soybean plants and soil. Proc. NEWCC 20:619-626.
7. CRAFTS, A. S. and S. YAMAGUTCHI.1964. The autoradiography
of plant materials. California Agr. Expt. Sta. Manual 35.
143 p.
8. CROSBY, D. G. 1964. Metabolites of 2,4-dichlorophenoxyacetic
acid (2,4-D) in bean plants. J. Agr. and Food Chem. 12:3-6.
9. CURTIs, D. F. and D. G. CLARK. 1950. An introduction to plant
physiology. p. 385. McGraw-Hill, New York.
10. FANG, S. C. 1958. Translocation and metabolism of 2,4-D-1-
C"4in pea and tomato plants. Weeds 6:178-186.
11. HAIJGAARD, G. and L. TUMERMAN. 1956. Reaction of al(loses
with amino compounds. Arch. Biochem. Biophys. 65:86-92.
12. SUTHERLAND, M. L. 1961. Amiben-C'4 preemergent tracer study
in soybeans. Technical report. Amchem Products, Inc.,
Ambler, Pa. 22 p.
13. SWAN, D. G. and F. WV.SLIFE. 1965. The absorption, translo-
cation, and fate of amiben in soybeans. Weeds 13:133-138.
14. SWANSON, C. R. 1965. Metabolic fate of herbicides in plants.
U.S.D.A., ARS publication 34-36. 36 p.
15. VANSLYKE, D. D., J. PLAZIN, and J. R. WEISINGER. 1951.
Reagents for the VanSlyke-Folch wet carbon combustion.
J. Biol. Chem. 191:299-304.
HARRISON2
be of several age classes so that only a part is at the most
susceptible age for control at any given time. The
presence of dormant seeds in agricultural soils provides
the weed with a reserve that may restore its population
after several years. Seed dormancy presents a real barrier
to the exploitation of modern chemical methods for the
control of weeds (7).
This paper deals with some aspects of the germination
and dormancy in wild carrot. Wild carrot, which has been
classified as a noxious weed by the Ontario Weed Control
Act, 1960, has been found in these studies to be an an-
nual, biennial, or short-lived perennial, propagating by
seed. Since attempts to propagate wild carrot from
cuttings of the fleshy storage organ have proved unsuccess-
ful, the maintenance of the population of this weed must
depend solely on the successful germination of seed.
In wild carrot, each fruit develops from a flower in
one of the compound umbels (or flower clusters). The
fruit is schizocarp, composed of two mericarps which are
held together on their flattened faces. Each mericarp
(half-fruit) has five ribs with four rows of large spines
(Figure 1) and contains a single seed with a small embryo
and abundant endosperm. Mericarps are referred to in
this paper as seeds.
Seeds of the Umbelliferae are noted for their dormancy
and poor germination due to immature embryos and
embryo-less seeds, respectively (3). Sylwester (8) found
that not all seeds germinate the first year, but some may
be (lormant up to 7 years.
It has been statedl that fresh see(ls of cultivated carrot
(D. carota var. sativuts (Hoffm.)) germinate better when

Wiv
Figure 1. Wild carrot seed: (a) outer curved surface with rows of
spines and ridges; (b) surface of seed with fruit coat removed;
(c), (d), and (e) sections showing relative size of seed coat, endo-
sperm, and embryo.
exposed to light; the seeds are receptive to the light after
having been soaked (1, 2). However, the light require-
ment becomes less important during dry storage (9).
Germination in the dark is increased with increased oxy-
gen concentration (4).
MATERIALS AND METHODS
Field observations and experimental studies were con-
tiucted in abandoned pastures in Wellington County, 40
miles north of Guelph. In one study, the location of indi-
vidual plants was mapped several times during
sive growing seasons. Two soil textures, sandy and clay
loams, in adjacent areas were used. Ten permanent
quadrats (50 x 50 cm) of each texture were mapped, using
characteristic symbols for each time of mapping.
quadrats on soil of each texture were in areas of low plant
density and five in areas of high density.
Seeds were collected from 10 areas at various times
during each season. Some seeds were tested for immediate
germination, others stored in paper bags at room temper-
ature and at 6 C, and some seeds were stored under water
at 6 C. Testing was conducted on seed untreated, or
treated in one of the following ways: with and without
seed coat, with light and heavy mechanical scarification,
soaked and chilled for several days, kept in the dark,
pricked in the endosperm, or with pieces of endosperm
removed.
Germination was tested on weighed lots of 100 seeds.
Hundreds of lots were put on 9 cm diameter (thick) filter
paper moistened with 4 ml of distilled water in petri
dishes. The petri dishes, in stacks of 7, were placed in
plastic bags or wrapped in new aluminum foil and set
on moist sand in large (20 liter) jars. The jars were
immersed to the neck in water of a constant temperature
bath at 16, 19, 21, and 25 C and given a continuous
light or a 16-hr photoperiod from a 100 watt bulb at
3gt. Germination also was tested in aerateto water or
pCosphate buffer solutions in Kolle flasks, in petri dishes
with expanded mica, soil, or sand, and in flower pots at
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W E E D S
several depths of soil. A few lots of 50 seeds in smaller
petri dishes were tested with or without an alternating
pressure. A siphoning device caused a pulse of 1/2 atmos-
phere above normal each 5-min cycle to be transmitted to
a glass jar containing these smaller samples.
RESULTS AND DISCUSSION
Field observations demonstrated the non-uniform size
and appearance of seedlings in all plots. Germination
occurred mainly in the spring, followed by successive
flushes in summer and early fall. The soaking of the
soil by heavy rains influenced the time of germination
and the number of seeds germinating. The first seedlings
to emerge in the spring were on sandy loam. Populations
of similar size range developed on both soil textures with
as many as 68 non-flowering plants per quadrat from
June to September. During the winter, the populations
were severely reduced, even to extinction on some quad-
rats. In June, the largest population was present on sandy
loam quadrats; however, the lower mortality and the
higher summer germination increased the population
on clay loam. There was considerable difference between
replicates. The population on one clay loam quadrat
dropped from 46 to 25 seedlings between July and
September, while on a replicate the population increased
from 4 to 15.
After-ripening occurred in wild carrot seeds. The
germination level in fresh seed was low and increased
with time for any given collection of seeds. Table 1 gives
Table 1. Germination expressed as percent of 5 lots of 100 seeds.
Seeds collected September 26, 1963, and stored at 6 C in paper
bags.
2 succes- Oct. 4 Dec. 6 Jan. 27 Mar. 27
Seed source 1963 1963 1964 1965
.1...
2.0.............
.......
O
6
19
7
22
19
22
3...1 .......1 1 1 5
Five
the average percent germination of five replicates of
seed. Viable seeds gave increased germination with time,
which could be attributed to some after-ripening process.
Even one year after collection, some seeds did not germin-
ate although they had live embryos which appeared
mature. These seeds still had the potential to develop into
plants.
After-ripening normally occurs in the field as seeds
over-winter after having been dispersed in the late sum-
mer. However, seed which remains attached to the plant
may be so immature that ripening does not take place.
A sample of these seeds was examined. Frequently, the
seeds of wild carrot in the fields are not dispersed until
late in the winter. It is possible that these seeds may be
blown some distance across the snow. On February 19,
1963, seeds were collected, hand rubbed, and tested for
germination. Only three seeds of 1,200 germinated. How-
ever, wind-blown seeds in the winter, even with ex-
tremely low germination, could invade new areas at a
distance.
The correlation between seed size and germination was
found to be significant only in some cases. Two hundred
and fifty lots were germinated under standard conditions
202
 DALE AND HARRISON:
in the laboratory. The lightest lot to germinate averaged
0.5 mg per seed, and the heaviest 1.8 mg. Germination
was below 40%ofor all lots of seeds with an average weight
of less than 0.7 mg per seed, but there was no other cor-
relation between seed weight and percent germination.
Seed weight increased from the center of each flower
cluster to the outside, and also varied from cluster to
cluster and plant to plant. Seed lots from individual
flower clusters were germinated separately, but in only
one case was correlation apparent between seed weight
and percent germination. This occurred with 2,500 seeds
from the one flower cluster of highest germination.
Weight per lot from this plant was 100.9 mg to 77.2 mg
andIgermination from 89%oto 73%. The mean weight of
100 seeds was 91.7 mg and the average germination was
79%o.For the seed lots from this flower cluster, there
was correlation between germination and weight of seed.
In Table 2, data are presented showing ranges of seed
number, weight, and germination from flower clusters
Table 2. Seeds from selected flower clusters, giving size of sample
used, mean weight of 100 seeds, and percent germination.
Number of seeds.. 700 500 1000 800
Weight, mg ...... 106 100 92 88
Germination %... 19 8 34 35
collected at one time in a single field and tested after
one year of storage in paper bags at room temperature.
The most striking feature is the diversity of the data.
No evidence was apparent of any pattern or relationship
between the number of seeds per flower cluster, lot
weight, and germination.
Temperature increased the speed as well as percent of
germination. During the first week, 80% of the final
germination occurred at higher temperatures, but only
50% at the lower temperatures. Table 3 presents data
Table 3. Germination, expressed as percent, of 3 lots of 100 seeds
each at 16-hr photoperiod and in darkness at 4 temperatures.
Temp.
C
Light
treatment
Germination
percent
Temp.
C
16 16 hr 18 21
Dark 21
16 hr 20 25
19
Dark 28
for dark and light treatments at four temperatures. These
data show that the sensitivity of wild carrot seeds to
light is not a simple matter but is dependent upon the
temperature at the time of soaking and/or germination.
Tests were made with alternating temperature and cold
treating of soaked seeds. The effects of these treatments
were not apparent in any pattern.
Germination was significantly lower on wet mica than
on wet soil. This could be attributed to the difficulty of
transfer of water to the seed from an open-textured sub-
strate. Low germination also was found in field plots on
dry friable muck soil. Greenhouse studies using soils of
different textures confirmed the findings from the field
plots that seeds were slow in soaking up water in coarse
textured material.
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WILD CARROT SEEDS
Seeds of wild carrot are roughly hemispherical with
a flat side, the curved surfaces being covered with spines
(Figure 1). The position of the spines, up or down, when
the seeds are on filter paper is reported by Cavers3 to
cause a significant difference in germination. However,
tests of 900 seeds showed this not to be critical.
When seeds were planted in loam soil in flower pots at
depthsof 14, 1,2, 1, 2, and 4 in, those at .1/ in gave highest
germination. Seeds germinated from all levels, but not all
seedlings developing from 2 and 4 in reached the surface.
Although it may be correctly argued that most weed seeds
receive conditions favourable for germination near the
soil surface (7), these data on the effect of light and depth
in soil on germination of seeds of wild carrot support
the observation that most forms of dormancy are broken
here, rather than supporting a thesis that light sensitivity
is of importance.
Seeds were stored underwater at 6 C for periods of
1, 3, and 6 weeks. Germination then was tested on filter
paper in petri dishes and also in aerated distilled water
in Kolle culture flasks. Soaking in water for 6 weeks or
less did not adversely affect germination. Subsequent
germination was the same in aerated distilled water and
1300 1900 1000 on filter paper.
84 71 68
24 11 15 Phosphate-citrate, phosphate-borate, and potassium-
sodium-phosphate buffers were used to determine
whether pH influenced the germination in 60 lots of
seed. On filter paper, the pH range used was 3.0 to 8.0,
and solutions from 5.6 to 9.2. At each pH rating, percent
germination differed sufficiently between replicates to
include the means of percent germination for any other
pH. The pH, per se, had no effect on germination.
In tests made in 1962, stacks of petri dishes without
plastic bags were placed directly into large jars contain-
ing moist sand. Invariably, a gradient in percent germin-
ation was found, with less germination in the upper
dishes. More uniform and higher germination resulted
when the dishes were first stacked in plastic bags closed
with a rubber band and then placed in the jar. Apparent-
ly, a more uniform and moist environment surrounding
the seeds was critical, whereas the reduction in the avail-
Light
treatment
Germination
percent
ability of oxygen from the air was not significant.
When seeds were tested in petri dishes, germination
16 hr 34
Dark 33 was enhanced by some severe treatments with sandpaper.
16 hr 35
Dark 26 However, other treatments, milder and more severe, or
pricking, did not improve the rate of germination.
Methods of treatment favorable to germination response,
resulted in seeds which had polished and rubbed surfaces,
with parts of the seed coats and endosperm removed.
Examination of the fruit structure revealed that the
embryo was completely embedded in endosperm (Figure
1). When the small cap of endosperm covering the emerg-
ing tip of the radicle was removed, germination under
standard conditions was higher. Tests of 500 treated seeds
showed that 55% germinated when the endosperm cap
was removed, whereas only 28%oof the untreated controls
germinated. A mechanical impedance imposed by a layer
of endosperm over the end of the embryo is responsible
for some seeds failing to germinate.
A siphoning device was used to test the effect of alter-
nating pressure on the impedance presented by the endo-
3Personal communication, 1964.
203
 W E
sperm. Two replicates of 10 lots of 50 seeds each, on
filter paper in small petri dishes wetted with 2 ml of
distilled water, were placed in the flask under continuous
light for 10 days. After that period, germination was
determined and ungerminated seeds were examined
under the microscope. Of the original 1,000 seeds, 400
had germinated and 300 had embryos but were as yet
ungerminate(l. These seeds were treated by removing the
small cap of endosperm that covered the radicle, and then
were returned to petri dishes for a further period of 10
days under standard conditions. With the removal of this
restriction of endosperm, half the seeds treated, or 15%
of the original number, then germinated. However, the
other 150 treated seeds with embryos (also 15%oof the
original), remained which did not germinate in spite of
the removal of the cap of endosperm. The effect of the
siphoning apparatus was to change the stress on the endo-
sperm cap so that some embryos could overcome the
mechanical impedance with artificial alternating pres-
sure similar to a squeezing action. This might have
simulated conditions of freezing and thawing. Other
embryos must have the cap removed to germinate im-
mediately. Discounting embryoless or rotted seeds, alter-
nating pressure and endosperm cap removal produced
80%cof the initial potential for germination.
The pattern of germination of wild carrot and other
weed seeds differs from the remarkably uniform be-
haviour of seed of crop plants. In wild carrot, not only
does this variation occur between seed lots collected from
plants in a single field at one time but also between seeds
from umbels on the same plant and, less significantly,
between seed lots from the same umbel. The germination
of a seed depends upon the state of the seed (whether
dormant or not) and the environment surrounding the
seed.
Germination of wild carrot is influenced by tempera-
ture but is most frequently limited by the amount of
water reaching the seed and being absorbed by it.
Harper (6) has pointed out that the surface of the soil
with its intricate patterns of different types of microsite,
offers to this and to many other weed seeds an oppor-
tunity for germination which may be denied the seed
when it is buried under the soil.
Dormancy in weed seeds has been classified into the
three categories of innate, induced, and enforced (7). In-
nate (lormancy is a property of ripe seed as it leaves the
p)lant. In wild carrot, innate dormancy is due to the
204
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E D S
mechanical barrier of the endosperm which, normally, is
digested at the tip of the radicle, allowing the embryo
to grow. In enforced dormancy, the environment prevents
germination, for example, when seed is buried in the
course of ploughing. Field observations show this condi-
tion to be frequent in wild carrot.
Control of plants such as wild carrot, which propagate
solely by seed, should be aimed at the reproductive phase
which is most vulnerable. However, control of this weed
is complicated by the fact that germination has been
found to occur in several flushes, each associated with
a rain that soaked the soil, following the breakdown of
the mechanical impedance of the endosperm. The popula-
tion of seedlings is not of uniform age, making chemical
control of seedlings crude. Control should be aimed at
the prevention of seed formation.
ACKNOWLEDGMENTS
The authors wish to thank: the Crop Science Depart-
ment, University of Guelph, for assistance in problems of
cleaning, counting, and scarification; Mr. P. Tyshuk for
conducting many germination tests; and the wife of the
senior author for cleaning, counting, and weighing many
thousands of seeds.
LITERATURE CITED
1. CROCKER,
W. 1936. Effect of the visible spectrum upon the
germination of seeds and fruits, p. 791-828. In B. M. Dugger
(Ed.), Biological Effects of Radiation, Vol. 2. McGraw-Hill
Book Co., New York.
2. EVENARI,M. 1956. Seed germination, p. 519-549. In A. Hol-
lander, (Ed.), Radiation Biology, Vol. 3. McGraw-Hill Book
Co., New York.
3. FLEMION, F. and E. T. HENDRICKSON. 1949. Further studies on
the occurrence of embryoless seeds and immature embryos in
Umbelliferae. Proc. Boyce-Thompson Inst. 15:291-297.
4. GARDNER, W. A. 1921. Effect of light on the germination of
light-sensitive seeds. Botan. Gaz. 71:249-288.
5. HARPER, J. L. and G. R. SAGAR. 1953. Buttercups-some aspects
of the ecology of buttercups in permanent grasslands. Proc.
Br. Weed Cont. Conf.:256-265.
6. . 1959. Factors controlling plant numbers, p. 119-132.
In Harper, J. L., (Ed.), The Biology of Weeds. Blackwell,
Oxford.
7. . 1959. The ecological significance of dormancy and
its significance in weed control. Proc. Internat. Conf. for
Crop Protection, Hamburg 4:415-420.
8. SYLWESTER, E. P. 1960. Beware of wild carrot. Hoard's Dairy-
man 105:330-331.
9. TOOLE,E. H., S. B. HENDRICKS,H. A. BORTHWICK, and V. K.
TOOLE. 1956. Physiology of seed germination. Ann. Rev.
Plant Physiol. 7:299-324.