Embryo and Seed Coat Factors Produce Seed Dormancy in Capeweed (Arctotheca Calendula)

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Embryo and seed coat factors produce seed dormancy in capeweed (Arctotheca
calendula)

Article  in  Crop and Pasture Science · October 2000


DOI: 10.1071/AR00050

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C S I R O P U B L I S H I N G

Australian
Journal of
Agricultural
Research
Volume 51, 2000
© CSIRO 2000

A journal for the publication of original contributions


towards the understanding of an agricultural system

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the Australian Academy of Science
Aust. J. Agric. Res., 2000, 51, 849–54

Embryo and seed coat factors produce seed dormancy in


capeweed (Arctotheca calendula)

A. J. ElleryA and R. Chapman

CSIRO Plant Industry, Private Bag 5, Wembley, WA 6913, Australia.


A
Corresponding author; email: a.ellery@ccmar.csiro.au

Abstract. Capeweed [Arctotheca calendula (L.) Levyns.] is common in annual pastures of southern Australia. The
physiological basis of seed dormancy in capeweed seeds was studied to determine the likely role of dormancy in
the success of this species. Two dormancy mechanisms were identified: one embryo-based and the other imposed
on the embryo by the seed coat. Embryo dormancy could be overcome by leaching, suggesting that this form of dor-
mancy was caused by a water-soluble germination inhibitor, possibly localised in the cotyledons. Following an
initial after-ripening period, embryo dormancy was rapidly relieved under conditions experienced on the soil
surface during summer. Ungerminated embryos entered secondary dormancy in winter. Seed coat-imposed dor-
mancy persisted longer than primary embryo dormancy, but was not associated with mechanical constriction of
embryo growth, or with limited water or oxygen supply to the embryo.
Embryo and seed coat dormancy mechanisms interacted to maintain a high level of dormancy in these seeds at
all times. Thus, a large proportion of seeds produced might enter a persistent seed bank, enabling re-establishment
following years when seed production is limited by unfavourable seasonal conditions or cropping activities.

Additional keywords: seed bank persistence, false break, germination, embryo dormancy.

Introduction matic zone are generally hot and dry (Buddenhagen 1990),
Capeweed [Arctotheca calendula (L.) Levyns.] is an these conditions can be interrupted by out-of-season rain
extremely successful pasture plant in the Mediterranean cli- (Heady 1958). Summer rainfall events which stimulate ger-
matic zone of southern Western Australia. This composite mination, but do not sustain seedling development, are
annual was accidentally introduced from South Africa known as false breaks of season (Rossiter 1966). False
around 1830 and has become widespread throughout south- breaks appear to be particularly common in the Western
ern Australia. In low rainfall areas of Western Australia, Australian wheatbelt during March and April (Cornish 1985;
capeweed regularly dominates pastures (Arnold et al. 1985), R. Chapman and S. Asseng, unpubl. data).
out-competing more agronomically useful species. The sig- In species with short-lived seed dormancy such as subter-
nificance of capeweed as a weed of cropping is likely to ranean clover (Trifolium subterraneum L.), in which seed
increase if herbicide resistance becomes widespread in this softening is typically complete by late summer (Taylor
species (Lemerle et al. 1996). 1984), false breaks can cause germination of a large propor-
The dominance of capeweed in pastures has previously tion of the seed bank population, leaving an inadequate seed
been attributed to superior ability of its seedlings to tolerate pool for more favourable establishment periods. Dormancy
early season drought (Rossiter 1966; Hallett 1971). Recently mechanisms that may protect seeds from false breaks
though, it has been shown that mortality of capeweed include: seed coat impermeability to water, exclusion of
seedlings is high under the severe drought stress which is oxygen from the embryo, mechanical restriction of embryo
associated with erratic summer or early autumn rainfall expansion, and physiological blocks to germination within
(Thomson et al. 1998). the embryo. Although Hallett (1971) has shown that seed
An alternative explanation may be that capeweed’s seed coat impermeability to water is not involved in regulating
dormancy mechanism is better adapted than those of other dormancy in capeweed, other dormancy mechanisms have
species to the summer and early autumn rainfall distribution not been extensively studied in this species. Therefore,
of the region. Although summers in the Mediterranean cli- experiments were conducted to evaluate the relative impor-
© CSIRO 2000 10.1071/AR00050 0004-9409/00/070849
850 A. J. Ellery and R. Chapman

tance of embryo- and seed coat-based dormancy mecha- Experiment 3: seed coat treatments
nisms and to identify how these mechanisms might con- Seed coat-imposed dormancy was further investigated in this experi-
tribute to the success of capeweed in a Mediterranean-type ment, in which seeds were given 1 of 3 seed coat treatments: (i) seed
environment. coat left intact (intact); (ii) seed coat removed completely to expose the
embryo (removed); and (iii) seed coat ruptured. In the ruptured treat-
ment the tip and sides of the seed coat were removed with a scalpel
Materials and methods blade, to release any physical constraint of the seed coat on the embryo,
Three experiments were conducted to investigate the seasonal dynam- while maintaining the body of the seed coat around the embryo. These
ics of dormancy in capeweed seeds, and to identify the physiological treatments were replicated 4 times and were applied to 12-month-old
processes controlling germination. Capeweed seeds were obtained from seeds which had been stored in the dark in the air-conditioned labora-
a long-term crop rotation experiment at Jennacubbine (31°27´S, tory. Each replicate contained 20 seeds.
116°43´E), Western Australia. Rainfall records for this site are only
Germination tests
available for 1996–1999, and average annual rainfall over that period
was 353 mm. In all experiments, germinability of seeds or isolated Seeds were placed in 9-cm Petri dishes lined with 2 discs of filter paper
embryos was used as a measure of dormancy. (Macherey-Nagel 615), moistened with 5 mL deionised water. Petri
dishes were sealed with Parafilm to minimise evaporation. Germination
tests were conducted for 14 days in growth cabinets maintained at a
Experiment 1: seasonal changes in dormancy status day/night temperature regime of 25°C/17°C with a 12-h photoperiod
Seeds were maintained under constant temperature (22°C) or exposed (349 mmol/m2.s of PAR). This temperature regime was chosen because
to prevailing conditions on the soil surface between summer and the fol- it approximates soil temperature after out-of-season summer rainfall
lowing spring, to indicate whether changes in germinability were asso- (Taylor 1996). Several previous studies have indicated that germination
ciated with daily and seasonal fluctuations in soil surface temperature. of capeweed seeds is relatively insensitive to temperature within the
Intact, freshly shed capeweed seeds were collected from the soil surface range 15–30°C (Piggin et al. 1973; Bolger et al. 1996; Dunbabin and
on 15 October 1997 and stored in the dark at room temperature. Seeds Cocks 1999); therefore, the germination temperature is unlikely to have
were placed in nylon mesh envelopes and allocated to either of 2 treat- substantially affected germination percentage.
ments. In the first treatment (soil surface storage), seed envelopes were Numbers of germinated seeds were counted daily, and at the con-
placed on soil contained in plastic trays on 24 November 1997. The clusion of the germination test, viability of ungerminated seeds was
trays were placed on the soil surface underneath a transparent plastic assessed by tetrazolium assay (International Seed Testing Association
rain shelter, at CSIRO Floreat Park (31°55´S 115°50´E). In the second 1976). In this assay, bisected seeds and embryos were incubated in 1%
treatment (laboratory storage), envelopes were stored in darkness in an (w/v) 2,3,5-triphenyltetrazolium chloride (Sigma) (pH 7) for 3 h in the
air-conditioned laboratory at approximately 22°C. dark at room temperature, before examination. Germinability was
Germinability of both intact seeds and excised embryos was tested expressed as a percentage of viable seeds. For intact seeds, germination
at the time of collection and again 2 weeks after seeds were transferred was defined as visible penetration of the seed coat by the radicle,
to storage treatments. At 4-weekly intervals thereafter, until May 1998, whereas for embryos and embryonic axes it was defined as extension
4 replicate seed envelopes were retrieved from each of the 2 storage and positive geotropism of the radicle.
locations and germinability of whole seeds and embryos was measured.
From May 1998 until October 1998, whole seeds continued to be tested Excision of embryos
monthly and embryos every 2 months. In each germination test, 50 Embryos were excised from mature capeweed seeds after soaking seeds
intact seeds and 20 embryos per replicate were used. in water for about 30 min to soften the seed coats. The tip and sides of
Soil surface temperature under the rain shelter was recorded the seed coat were then removed with a scalpel blade. Light pressure
throughout the experiment at 30-min intervals using thermocouples was applied to the sides of the seed with forceps and the scalpel blade
connected to a data logger (Datataker 50, Hinco). Temperatures was used to separate the halves of the seed coat. Embryos were removed
observed under the rain shelter were compared with those expected from the adhering seed coat half and placed directly on to moist filter
from measurement of temperature on bare soil (Taylor and Revell paper in Petri dishes. In all experiments the term ‘intact seeds’ is used
1999). to refer to seeds whose seed coat was not removed, and the term
‘embryos’ refers to seeds from which the seed coat was removed.
Experiment 2: embryo treatments
Statistical analysis
The objective of this experiment was to investigate the physiological
Germination percentage data were arcsine transformed prior to analysis to
basis of embryo dormancy. The effects of leaching and cotyledon
correct for the non-linear distribution of variance associated with propor-
removal on the germinability of embryos and intact seeds were mea-
tions (Mead and Curnow 1983). The effects of interactions between time
sured in January 1998, using seeds collected in October 1997 and stored
and storage environment on germinability of intact seeds and embryos
in darkness in the air-conditioned laboratory. Each of 3 treatments
were determined by factorial analysis of variance. Pair-wise comparisons
(control, leached, and bisected) was applied to seeds with and without
between treatments were made using least significant difference at
seed coats and replicated 4 times. Control seeds and embryos were
P = 0.05. The effects of leaching, cotyledon removal, and seed coat manip-
imbibed and germinated, using 20 seeds and 20 embryos per replicate.
ulation were likewise determined in separate factorial analyses of vari-
In the leached treatment, each replicate containing 20 intact seeds or
ance. Untransformed percentage data are presented in graphical form.
embryos was placed in a Buchner funnel lined with filter paper
(Whatman No. 42). A peristaltic pump was used to rinse seeds and
embryos with approximately 1 L of deionised water over a 24-h period.
Results
In the bisected treatment, replicates of 20 intact seeds and 20 embryos Experiment 1: seasonal dormancy of capeweed seeds
were cut transversely to remove the cotyledons. This generated embry-
onic axes with the seed coat either present or absent. Germinability of
Soil temperatures
control seeds and embryos, leached seeds and embryos, and embryonic Soil surface temperatures exhibited a diurnal cycle with
axes was then assessed in a germination test (described below). extremes of approximately 60°C and 20°C in summer,
Seed dormancy in capeweed 851

Fig. 2. Expt 2: effect of leaching and cotyledon removal on the ger-


minability of intact seeds and embryos. Bars topped by the same letter
are not significantly different (P > 0.05).

Germinability of intact seeds stored at room temperature


did not differ significantly from zero at any time (Fig. 1b). In
contrast, an increase in germinability of seeds stored on the
soil surface was observed in late February, at which time
seeds had been exposed to a daily temperature fluctuation of
approximately 20°C to 60°C for about 2 months.
Germinability reached a maximum of approximately 14% of
viable seeds by April (Fig. 1b) and remained at this level
throughout May. A downward trend in germinability was
observed throughout the winter months but the decline from
maximum germinability was not significant.
Most embryos from seeds in both storage treatments also
remained dormant through December but germinability
increased in both treatments from January 1998 (Fig. 1c).
Fig. 1. Expt 1: (a) average monthly maximum (䉭) and minimum (䉱) Germinability of embryos from seeds held at room tempera-
temperatures (°C) measured at the soil surface with thermocouples; ture increased at a near-linear rate, reaching approximately
changes in germinability of (b) intact seeds and (c) embryos dissected
from seeds stored in the laboratory (䊉) or on the soil surface (䊊) from
60% of viable embryos by September. In contrast, embryos
late November until September and germinated monthly. Dates marked excised from seeds stored on the soil surface showed a
on the X-axes represent the first of each month. Values marked with the marked seasonality in germination response. Dormancy was
same letters are not significantly different (P > 0.05). relieved rapidly during mid- to late summer, so that
maximum germinability (about 70%) was obtained by
falling to approximately 35°C and 10°C in winter months March 1998, when the average daily soil temperature was
(Fig. 1a). The average daily temperature at the soil surface approximately 35°C. Germinability remained high through-
ranged from 42.1°C in January to 19.6°C in July. The mag- out autumn but fell significantly between May and July. This
nitude of daily temperature fluctuation fell from about 40°C reduction in germinability was associated with a drop in
in summer to about 25°C in winter. The temperatures mea- average daily temperature <25°C.
sured were consistent with those previously measured on
bare soil. Experiment 2: embryo dormancy
Intact seeds did not germinate following either imbibition in
Seed germinability Petri dishes (control) or leaching for 24 h (leached) (Fig. 2).
Immediately after seed collection, neither embryos nor Germinability of control embryos was 21.4%, whereas that
whole seeds germinated (Fig. 1b, c); however, seed viability of leached embryos was 73.8%. Bisection of seeds and
was in excess of 90% (data not shown). Thereafter, the embryos to remove cotyledons also promoted germination of
responses of embryos and intact seeds to storage treatments embryonic axes compared with intact seeds and embryos.
differed substantially. Germinability of seeds with seed coats present increased to
852 A. J. Ellery and R. Chapman

In the case of capeweed, the seed coat did not appear to


impose a physical barrier to germination because there was
no significant response to removal of mechanical constraints
on embryo growth (by the rupturing treatment) in Expt 3.
Free diffusion of water and oxygen to the embryo were also
possible in this experiment, indicating that the seed coat did
not inhibit either of these processes. It has previously been
shown that scarification of capeweed seeds does not enhance
germination (Hallett 1971), further indicating that water
uptake is not restricted by the seed coat. In addition to these
results from intact seeds, results from Expt 2 with bisected
seeds also indicated that the seed coat did not restrict oxygen
uptake. Germination of bisected axes, thus exposed to atmo-
spheric oxygen but still in contact with the seed coat, was
significantly lower than germination of axes from which
Fig. 3. Expt 3: effects of rupturing and removing seed coats on ger- seed coats had been removed. This indicated that the seed
minability. Bars topped by the same letter are not significantly different coat inhibited germination even when oxygen was not limit-
(P > 0.05). ing. The possible presence of a germination inhibitor in the
seed coat has not been evaluated.
10.3% when cotyledons were removed. When both the seed Embryo dormancy
coat and cotyledons were removed, germinability of embry-
Enhanced germination of excised embryos in Expt 1 and the
onic axes increased to 82.5%. There was no significant dif-
response to leaching and cotyledon removal in Expt 2 indi-
ference between the germinability of these embryonic axes
cated the presence of an additional dormancy mechanism
and that of whole, leached embryos.
which was confined to the embryo.
Experiment 1 showed that embryo dormancy was present
Experiment 3: seed coat-imposed dormancy
at seed maturity in spring and at least 6 weeks of dry storage
Germinability of intact, 12-month-old seeds was 1.3% were required before it began to decline. After this initial
(Fig. 3). Rupturing seed coats to remove any physical con- period, the rate of dormancy decline was accelerated by
straints to germination did not significantly alter germinabil- exposure of seeds to conditions on the soil surface. The two
ity. Complete removal of the seed coat significantly most obvious differences between the field and laboratory
increased germinability to 77% of viable embryos. conditions to which the seeds were subjected were tempera-
ture (high and widely fluctuating at the soil surface) and
Discussion light. Exposure to light during dry storage has been reported
Two dormancy mechanisms appear to control the germina- to increase germinability of intact capeweed seeds
tion of capeweed seeds. The net effect of these seed coat- and (Chaharsoghi and Jacobs 1998). Although Quinlivan (1972)
embryo-based mechanisms in the present study was to speculated that dormancy relief in capeweed seeds was
produce a persistent dormancy in a large proportion of the related to high temperature, Dunbabin and Cocks (1999)
seed pool, and to control the seasonality of dormancy release have recently shown that seeds maintained in a dark oven
in those seeds that did become germinable. with a daily temperature fluctuation of 15°C–60°C, which
simulates the daily cycle of soil surface temperatures, did not
Seed coat dormancy lose dormancy as quickly as seeds placed on the soil surface.
The marked effect of seed coat removal on germinability in These findings indicate that both light and soil surface tem-
all 3 experiments indicated that some form of seed coat- perature may be factors in the relief of dormancy in cape-
imposed dormancy was involved in germination regulation. weed seeds, although their specific role in relief of embryo
Seed coat-imposed dormancy has been observed in many dormancy is currently unknown.
other species and may take a number of forms including Approximately 20% of the embryos in seeds stored on the
physical restriction of embryo expansion (Egley 1972; soil surface remained dormant during autumn. Dormancy
Junttila 1973; Watkins and Cantliffe 1983; Bradford 1990; release in these seeds may require a longer period of expo-
Welbaum et al. 1995) and water-impermeable seed coats sure to elevated and fluctuating temperatures than they
(Ballard 1973). Seed coats may also prevent uptake of received in this study. It is possible that these embryos would
oxygen by the embryo (Qi et al. 1993; Stabell et al. 1996), or become germinable if they were given additional exposure to
may contain germination-inhibiting compounds (Richmond summer soil surface temperatures, or exposure to a second
and Ghisalberti 1994; Plummer et al. 1995). summer in the soil. During winter, non-dormant seeds stored
Seed dormancy in capeweed 853

on the soil surface returned to a dormant state, but seeds exposed to late spring rainfall. The deep embryo dormancy
stored at 22°C did not, suggesting that the lower tempera- observed in freshly produced seeds would prevent germina-
tures experienced on the soil surface during winter played a tion during spring rainfall even if seed coat-imposed dor-
role in the development of secondary embryo dormancy. mancy failed to develop. Embryo dormancy, however, was
The large response to leaching obtained in Expt 2 indi- largely relieved by March when capeweed seeds were stored
cates that embryo dormancy may have been imposed by a on the soil surface; therefore, it is unlikely to provide a sig-
water-soluble germination inhibitor. The changes in ger- nificant barrier to germination during out-of-season rainfall.
minability of capeweed embryos over time observed in A previous study of seed bank dynamics in pasture commu-
Expt 1 may be related to a change in the level, activity, or sol- nities also showed that the largest increase in germinable
ubility of this putative germination inhibitor. Enhanced ger- capeweed seed population occurred between February and
mination following leaching or pre-incubation of embryos in March (Bolger et al. 1996), which is in close agreement with
water has been observed in a number of species (Bewley and the results observed for embryos in this study. Both results
Black 1982). These effects have been attributed to leaching indicate that seed dormancy provides good protection from
of germination inhibitors from the seed (Taylor and Rossiter false breaks which occur before March, but not from those
1967) or to increased sensitivity of embryos to endogenous occurring later in autumn.
gibberellic acids (Le Page-Degivry et al. 1990). The high level of seed coat-imposed dormancy observed in
Removal of the cotyledons from embryos in Expt 2 this study may provide additional benefits for the seed cohort
caused a similar germination response to leaching, suggest- by ensuring that seeds are maintained in a dormant state for
ing that the germination-inhibiting activity was localised to future years. Only 14% of viable seeds from this cohort
the cotyledons. Stimulation of germination in otherwise achieved germinability in the first year after seed production,
dormant embryos after removal of cotyledons or scutellum, even when exposed to potentially dormancy breaking condi-
a modified cotyledon in cereals, has been reported in a tions on the soil surface. Under field conditions the remainder
variety of other species (Bewley and Black 1982; Pinfield would have entered a persistent soil seed bank. These stored,
and Stutchbury 1990). Germination-inhibiting properties of dormant seeds may become germinable in future years, as
cotyledons have been attributed to their abscisic acid (ABA) observed by Dunbabin and Cocks (1999), enabling re-estab-
content. Removal of cotyledons, however, may stimulate a lishment of the capeweed population in years when seed pro-
wounding response involving evolution of ethylene or CO2, duction is limited by unfavourable environmental conditions
which may have stimulated germination (Leather et al. or cropping activities. The percentage of capeweed seeds
1992), rather than removal of ABA. entering the persistent seed bank may vary from year to year,
since dormancy can be strongly influenced by prevailing
Ecological significance of dormancy environmental conditions during seed development
We have shown that dormancy in capeweed involves pro- (reviewed by Fenner 1991). In order to estimate annual seed
cesses in both seed coat and embryo, and that embryo dor- bank inputs, it will be necessary to gain a more thorough
mancy shows a seasonal cycle. This pattern is typical for understanding of the mechanism underlying seed coat-
seeds of winter annual species with a physiological dor- imposed dormancy and its response to environmental condi-
mancy mechanism (Baskin and Baskin 1998). Dunbabin and tions. Nevertheless, seed coat-imposed dormancy appears to
Cocks (1999) have recently reported a seasonal dormancy be an effective means of distributing germination over a
cycle for capeweed, which was particularly marked in buried number of years.
seeds. In the present experiment, the interaction between
seed coat and embryos resulted in a high overall level of dor- Conclusions
mancy at all times. This work has identified dormancy mechanisms in both
The seasonal nature of dormancy in capeweed seeds may embryos and seed coats of capeweed seeds. Embryo dor-
benefit the population by restricting germination to the mancy displayed a marked seasonal cycle. Although this
period between autumn and early winter, when seedling sur- cycle may protect seeds from untimely germination in
vival is most likely. By avoiding germination either too early spring, summer, and winter, it did not appear to offer sub-
in the autumn, when false breaks are more likely, or too late stantial protection from the effects of false breaks occurring
in the winter, when plants are unlikely to survive due to com- in autumn. The mechanism of dormancy operating in the
petition with established plants, the dormancy pattern seed coat, however, appeared to be instrumental in maintain-
appears to optimise the distribution of germination within a ing the majority of seeds in a dormant state. This high level
single year. of residual dormancy can be expected to facilitate persis-
Embryo dormancy may afford capeweed seeds some pro- tence of a seed pool, which will aid the survival of capeweed
tection against rainfall after seed maturation. Capeweed populations across seasons of variable quality, and contribute
seeds, which develop earlier than other pasture species and to the success of capeweed in the Mediterranean environ-
may also be shed earlier (McIvor and Smith 1973), may be ment of Western Australia.
854 A. J. Ellery and R. Chapman

Acknowledgments Lemerle D, Yuan TH, Murray GM, Morris S (1996) Survey of weeds
and diseases in cereal crops in the southern wheat belt of New South
We thank Meg Flavelle for excellent technical assistance Wales. Australian Journal of Experimental Agriculture 36,
and Murray Siegert for provision of the Jennacubbine site 545–554.
through GRDC project DAW361. We also thank Matthew Le Page-Degivry M-T, Barthe P, Garello G (1990) Involvement of
Dunbabin, Clinton Revell, Robert Gallagher, and Graham endogenous abscisic acid in onset and release of Helianthus annuus
Taylor for useful discussions during preparation of the embryo dormancy. Plant Physiology 92, 1164–1168.
McIvor JG, Smith DF (1973) The effect of defoliation on seed produc-
manuscript. tion by capeweed (Arctotheca calendula). Australian Journal of
Experimental Agriculture 13, 676–680.
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