J.M. ZABALA, P.A. TOMAS, G.E. SCHRAUF AND J.A.

GIAVEDONI

Zabala, J.M., Tomas, P.A., Schrauf, G.E. and Giavedoni, J.A. (2009), Seed Sci. & Technol., 37, 248-251

Research Note
Seed dormancy in Elymus scabrifolius (Dll) J.H. Hunz.
J.M. ZABALA,1 P.A. TOMAS,1 G.E. SCHRAUF2 AND J.A. GIAVEDONI1
1

Facultad de Ciencias Agrarias, Universidad Nacional del Litoral, Kreder 2805, Esperanza (S3080HOF)
Argentina (E-mail: jmzabala@fca.unl.edu.ar)
2
Facultad de Agronoma, Universidad Nacional de Buenos Aires, Av. San Martn 4453 - Buenos Aires
(C1417DSE), Argentina

(Accepted August 2008)

Summary
The objective of this study was to determine the effect of different germination temperatures and dry storage
on the germinative behaviour of E. scabrifolius. Seeds of three lines from different populations, which showed
contrasting germinative behaviour in preliminary experiments, were incubated under four temperature regimes:
15C, 20C, 15-20C and 25C. Germination tests were performed with freshly harvested seeds and seeds kept
dry for nine months in paper bags at 25C. The results presented in this work suggest that in E. scabrifolius:
a) seeds do not have putative dormancy; b) seeds require low temperature for germination; c) germination is
prevented at 25C; d) alternating temperature was better than a constant 15C temperature regime for final
percentage germination.

Experimental and discussion

Elymus scabrifolius (Dll) J.H. Hunz is native to Argentina and Uruguay and is valued as
a forage resource (Parodi, 1940; Covas, 1978). The domestication of this species began
during the fifties in the twentieth century. Since then, only one cultivar has been released
but was not successfully widespread because of its poor germinative behaviour. Andres
(1986) considered that this species was in imminent danger of extinction. Based on these
assumptions, many plant collections have been conducted by the Forage Plant Breeding
Program of the Facultad de Ciencias Agrarias (Universidad Nacional del Litoral) during
the last ten years. There is little published information regarding the germinative behaviour
of E. scabrifolius. Guma and Alonso (1997) and Giavedoni (2000) found that alternating
temperatures promoted germination in one population and four lines respectively. Andrs
(1986) and Giavedoni (2000) observed that the percentage germination of viable seeds
varied among genotypes when incubated at constant temperature, and attributed this fact
to seed dormancy. Seed dormancy is a relative phenomenon which can be eliminated,
for example, during dry storage. This factor and a wide range of temperatures were not
analyzed in previous reports.
248

F O R MATT E D P R O O F

SEED DORMANCY IN E. SCABROFOLIUS

The objective of this study was to determine the effect of different germination
temperatures and dry storage on the germinative behaviour of E. scabrifolius.
Seeds of three lines from different populations, which showed contrasting germinative
behaviour in preliminary experiments, were incubated under four temperature regimes:
15C, 20C, 15-20C (16 hours and 8 hours respectively) and 25C. Germination tests
were performed with freshly harvested seeds and seeds kept dry for nine months in paper
bags at 25C.
Tests were conducted by placing 25 seeds in a Petri dish containing three Whatman
paper discs moistened with distilled water. Four replicate dishes were used for each line
and treatment. The number of germinated seeds was counted at 28 days (final percentage
germination) after sowing. Only seeds showing a 2 mm radicle length were considered
germinated. Viability of non-germinated seeds was determined at the end of the experiment
(AOSA, 1993).
The data were examined using INFOSTAT statistical package (INFOSTAT version
2006p.2). Prior to analysis percentage germination values were arcsin transformed;
however, untransformed means are also reported in the tables and figures. The final
percentage germination (FPG) of viable seeds was subjected to an analysis of variance
(ANOVA) to determine the significance of line, temperature treatment, storage and their
interactions. Differences between means values were tested for significance using a Tukey
test (p<0.05).
Variations in FPG were highly significant for line, treatment, storage and their
interaction (p<0.001 for all factors). The significant interaction terms of the analysis
reflect that the seeds from different lines gave rise to different germination responses
depending on the temperature treatment and storage time.
Temperature treatments were analyzed separately. ANOVA revealed significant line
storage time interactions at constant temperatures 20C and 25C. Freshly harvested
seeds of analyzed lines did not germinate at these constant temperatures, but lines 104
and 212 showed a significant increase in FPG after storage compared with line 912
(figure 1), and this line did not germinate at 25C after 9 months of storage. Differences
between alternating temperatures and constant 15C were significant when all lines were
analyzed together (92.8% vs. 77.5%, respectively). Although in these temperature regimes
differences in PFG between lines and time storage were not significant and lines showed
high FPG values.
The results presented in this work suggest that E. scabrifolius seeds do not have
putative dormancy but rather require low temperatures for germination. In this regard
alternating temperatures were better than a constant 15C temperature regime for final
percentage germination (FPG), while germination is prevented at 25C. There was a
progressive trend toward reduced germination at warm temperatures (20C and 25C).
This might be a genetic trait of the species, as in Allium sp. (Specht and Keller, 1997).
Our results were in agreement with other published data for related species, where
alternating temperatures are optimal for germination (Knipe, 1973; Toole, 1976; AOSA,
1993; Young et al., 2003). Often alternating temperatures in grass species are more
favourable for germination than constant temperatures, since in natural environment seeds
are exposed to alternating rather than constant temperatures (Simpson, 1990; Baskin and

F O R M A T T E D

P R O O F

249

J.M. ZABALA, P.A. TOMAS, G.E. SCHRAUF AND J.A. GIAVEDONI

15/20C

15C

100

FPG (%)

80
60
40
20
0

20C

100

25C

b b

104
212
912

FPG (%)

80
60
40

20
0

Freshly mature
seeds

9 months of
storage

Freshly mature
seeds

9 months of
storage

Figure 1. Final percent germination (FPG) for seeds of three lines exposed to different temperature treatments
with and without storage. Means with different letters are different at the 5% level according Tukeys test.

Baskin, 1998). The temperature fluctuation sensitivity some seeds show for germination
may act as an environmental sensor of surrounding vegetation (gaps), soil depth and/or
depth of submergence under water (Fenner and Thompson, 2005). On the other hand,
the variable germinative behaviour of the lines at 20C in freshly mature seeds would
indicate a mechanism of warm temperature dormancy in accordance with reports for
other temperate grass species (Poljakoff-Mayber et al., 1990, 2002; Black, 1991; Fandrich
and Mallory-Smith, 2005; Matus-Cadiz and Hucl, 2005.). In this regard seed dormancy
at warm temperature has been proposed by Andrs (1986) and Giavedoni (2000) as a
character present in E. scabrifolius seeds.
Future studies on the germination behaviour of this species should involve pretreatments or the application of compounds known to alleviate dormancy (cold treatment,
GAs, light, etc.) and study the influence of external structures that cover the caryopsis
on germination. These will determine the existence or not of seed dormancy at warm
temperatures and reveal the seed germination patterns in E. scabrifolius.

Acknowledgements
This study was supported by a research grant from Universidad Nacional del Litoral and
Agencia Nacional de Promocin Cientfica y Tecnolgica (Argentina). We thank Dr.
Jose Francisco Pensiero and Ing. Agr. Hugo Francisco Gutierrez for comments on the
manuscript.

250

F O R MATT E D P R O O F

SEED DORMANCY IN E. SCABROFOLIUS

References
Andres, A.N. (1986). Variabilidad gentica en poblaciones naturales de Agropyron scabrifolium (Doell) Parodi.
[Genetic variation in native populations of Agropyron scabrifolium (Doell) Parodi]. Tesis M.Sc. Pergamino,
INTA- Universidad Nacional de Rosario, 93 pp.
Association of Official Seed Analysts (AOSA). (1993). Rules for testing seed. Proceeding of the Association of
Official Seed Analyst. Journal of Seed Technology, 16, 1-78.
Baskin, C.C. and Baskin, J.M. (1998). Chapter 3. In Seeds. Ecology, biogeography, and evolution of dormancy
and germination, (Elsevier inc.), pp. 27-47, San Diego, Academic Press.
Black, M. (1991). Involvement of ABA in the physiology of developing and mature seeds. In Abscisic acid:
physiology and biochemistry (eds. W.I. Davies and H.G. Jones), pp. 99-109, Bios Scientific Publishers,
U.K.
Covas, G. (1978). Forrajeras indgenas: Especies que requieren un plan de conservacin de germoplasma. [Native
Forages: Species urging a germplasm conservation programme]. Ciencia e Investigacin, 34, 209-213
Fandrich, L. and Mallory-Smith, C. (2005). Temperature effects on jointed goatgrass (Aegilops cylindrica) seed
germination. Weed Science, 53, 594-599.
Fenner, M. and Thompson, K. (2005). The Ecology of Seeds. Cambridge Univ. Press 250 pp
Giavedoni, J. (2000). Evaluacin de poblaciones y progenies de agropiro criollo (Elymus breviaristatus subsp
scabrifolium) coleccionadas en el centro norte de la provincia de Santa Fe. [Evaluation of populations and
progenies of argentine wheatgrass [Elymus breviaristatus subsp. scabrifolium collected in north and centre
Santa Fe.] Tesis M.Sc. Pergamino, INTA- Universidad Nacional de Rosario.
Guma, I.R. and Alonso, S. (1997). Germination and seedling growth in Elytrigia scabrifolia and E. scabriglumis.
Seed Science and Technology, 25, 343-350.
Knipe, O.D. (1973). Western wheatgrass germination as related to temperature, light, and moisture stress. Journal
of Range Management, 26, 68-69.
Matus-Cadiz, M.A. and Hucl, P. (2005). Rapid and effective germination methods for overcoming seed dormancy
in annual canarygrass. Crop Science, 45, 1696-1703.
Parodi, L.R. (1940). Estudio crtico de las Gramneas austral-americanas del gnero Agropyron. [Critical study
about austral-american grasses of the genus Agropyron]. Revista del Museo de la Plata, 3, 63 pp.
Poljakoff-Mayber, A., Corbineau, F. and Cme, D. (1990). A possible mechanism of high temperature dormancy
regulation in seeds of Avena sativa L. (cv. Moyencourt). Plant Growth Regulation, 9, 147-156.
Poljakoff-Mayber, A., Popilevski, I., Belausov, E. and Ben-Tal, Y. (2002). Involvement of phytohormones in
germination of dormant and non-dormant oat (Avena sativa L.) seeds. Plant Growth Regulation, 37, 7-16.
Specht, C.E. and Keller, E.R.J. (1997). Temperature requeriment for seed germination in species of the genus
Allium sp. Genetic and Crop Evolution, 44, 509-517.
Toole V. K. (1976). Light and temperature control of germination in Agropyron smithii seeds. Plant and Cell
Physiology, 171, 1263-1272.
Young, J.A., Clements, D.C. and Jones, T. (2003). Germination of seeds of big and bottlebrush squirreltail.
Journal of Range Management, 56, 277-281.

F O R M A T T E D

P R O O F

251