VARIATION IN SEED GERMINATION BETWEEN ELYMUS SCABRIFOLIUS LINES FROM DIFFERENT HABITAT

Zabala, J.M., Tomas, P.A., Schrauf, G.E. and Giavedoni, J.A. (2009), Seed Sci. & Technol., 37, 245-250

Research Note
Variation in seed germination between Elymus scabrifolius
(Dll) J.H. Hunz. lines from different habitats
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
Elymus scabrifolius is a South American species, valued as a forage resource in areas with soil limitations. The
objective of this study was to determine the effect of different germination temperatures on the germinative
behaviour of twenty representative lines of E. scabrifolius from four different habitats. The results suggest that
in E. scabrifolius the germinative behaviour can be related with provenance. The observed pattern of variation
may be capitalized for future improvement programs and to determine new management rules for this species
when grown in native grassland or under sown pastures.

Experimental and discussion

Elymus scabrifolius (Dll) J.H. Hunz is a South American species valued as a forage in
salt-affected soils (Covas, 1978; Ranmuno, 1980; Vargas Lpez, 1982). The germinative
behaviour is a key characteristic for the domestication of this valuable forage grass. E.
scabrifolius does not have putative dormancy, however it does require low temperatures for
germination and warm temperatures could have an inhibitory effect on seed germination
(Andrs, 1986; Giavedoni, 2000; Guma y Alonso, 1997; Zabala et al., 2009).
The objective of this study was to determine the effect of different germination
temperatures on the germinative behaviour of lines of E. scabrifolius from different
habitats.
Twenty representative lines collected from four different populations (named 1, 2, 4
and 9) from the Argentine were used in this study. Populations 1 (lines 101, 102, 104, 105,
109, 119) and 2 (lines 201, 202, 204, 210, 212, 214) are located next to water courses of
sweet water (31 15 S, 60 37 W and 31 10 S, 60 25 W respectively) while populations
4 (lines 401, 402, 403, 414) and 9 (lines 901, 908, 912, 913) are located next to saline-sodic
water courses (29 31 S, 60 28 W and 30 55 S, 61 3 W respectively). In this report
245

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

the term lines makes reference to natural inbred plants (predominantly homozygous).
The lines were chosen at random from each population and seed multiplication was carried
out at the Facultad de Ciencias Agrarias (Universidad Nacional del Litoral), Santa Fe,
Argentina (31 25 S and 60 56 W).
Seeds of the twenty lines were incubated under four temperature regimes: 15C, 20C,
15-20C (16 hours and 8 hours respectively) and 25C. Seeds were harvested and kept
for nine months in freely permeable paper bags at about 25C until being used in the
experiments.
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 3, 5, 7, 10, 14, 21, 28
and 35 days 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 reported in tables and figures. The final percentage
germination (FPG) of viable seeds and the Coefficient of Germination (CG) estimated by
Maguire Index (Maguire, 1962) were subjected to an analysis of variance (ANOVA) to
determine the significance of line, temperature treatment and line temperature treatment
interaction. Differences between means values were tested for significance using a Tukey
test (p<0.05).
Germination started 7-10 days after sowing and showed the maximum percentage
germination after 28 days in almost all lines subjected to the temperature treatments 20C,
15-20C, and 25C. Germination was slightly delayed in seeds incubated at 15C, but the
percentage germination at 28 days was higher than 80% in all lines.
Variations in FPG and CG were highly significant between lines, treatments and for
their interaction (p<0.0001). Because of these significant interactions it was concluded
that the treatments had to be analyzed separately.
A significant variation for FPG and CG was found between the lines, with germination
being significantly higher and faster at 15C and 15-20C vs. 20C and 25C (figures 1
and 2). All the lines subjected to 15C and 15-20C treatments showed a FPG value
higher than 80%, and the highest differences in FPG between lines were found in the
20C treatment. In the 25C treatment a maximum of 16% germination was recorded for
all the lines.
When temperature treatments 15C and 15-20C were analyzed, there were significant
differences in FPG and CG values between lines and temperatures (p<0.001 for all factors).
In the 15-20C treatment, percentage germination and germination rate were higher than
15C (94.3% vs. 81.2% and 30.5 vs. 19.8 respectively).
We suggest, in agreement with Guma and Alonso (1997) that percentage germination
at 15-20C should be used in germination tests. We also showed that 10 and 28 days
would be useful as the initial and final germination dates respectively.
We also found some evidence for habitat correlation between lines and their
germination response. Guma and Alonso (1997) described geographical patterns in seed
246

VARIATION IN SEED GERMINATION BETWEEN ELYMUS SCABRIFOLIUS LINES FROM DIFFERENT HABITAT

100

15C

80
60
40
20
0

15/20C

100
80
60

FPG (%)

40
20
0

20C

100
80
60
40
20
0
100

25C

80
60
40
20
0
101 102 104 105 109 113 201 202 204 210 212 214 401 402 403 414 901 908 912 913

Lines
Figure 1. Final percentage germination (FPG) for seeds of lines exposed to different temperature treatments.

germination of two species of Elymus native from the central region of Argentina. In a
similar way, our results made it possible to determine the germination patterns related
to the origin of the lines. In all populations of E. scabrifolius that were tested the seeds
are dispersed in early summer and grow in areas characterized by intense rainfalls and
low evapo-transpiration during mid autumn and early spring. In mid autumn and early
spring the soil temperatures are similar to those that promote the germination in this study
(Panigatti, 1968; Cceres, 1980). Similar general germination patterns have been found
in other grass species such as Bromus tectorum L. and Elymus elymoides (Raf.) Swezey
(Beckstead et al., 1995).
247

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

45

15 C

30

15

0
45

15/20
C

30

15

CG

20C

45

30

15

0
45

25 C

30

15

0
101 102 104 105 109 113 201 202 204 210 212 214 401 402 403 414 901 908 912 913

Lines
Figure 2. Germination rate (CG) for seeds of lines exposed to different temperature treatments.

The variable behaviour between lines subjected to the 20C treatment, have been
associated to origin of the populations. Lines in populations termed "4" and "9" were
collected in a region known as Bajos Submeridionales, in Spartina spartinae tall
grasslands characterized by sodium-saline soils with poor vertical drainage and slow
surface drain. On the other hand, lines from populations designated as 1 (101, 102,
104, 105, 109, 119) and 2 (201, 202, 204, 210, 212, 214) were collected within an
area named Terraza anterior del Ro Paran, which has non-saline soils related to the
fluvial environment generated by rivers (Golln and Lachaga, 1939; Espino et al., 1983).
It is well known that some plant species, native to saline environments, are endowed
248

VARIATION IN SEED GERMINATION BETWEEN ELYMUS SCABRIFOLIUS LINES FROM DIFFERENT HABITAT

with mechanisms that facilitate their germination and establishment during a specific
season in which the harmful effects of salts are minimized (Sen and Mohammed, 1994).
The lines with lower germination capacity at constant temperatures (i.e. more specific
temperature requirement) were those collected in a saline region. Germination could
only occur during the wet season as mentioned above, when conditions are favourable
for plantlet growth in a humid environment with low salinity levels. Similarly in jointed
goatgrass (Aegilops cylindrica Host) analysis of seed germination showed that favourable
temperatures for germination coincide with temperatures of the wet seasons (Fandrich and
Mallory-Smith, 2005). Conversely, the lines native to non-saline origins, are capable of
germination under a wider range of temperatures. In this way the favourable period for
their germination extends throughout the year.
Both the observed pattern and the variation found between lines may be capitalized
for future improvement programs and may also be used to determine new management
rules for this species. Breeders select materials with a germinative behaviour that favours
a rapid and uniform seedling establishment from different sowing dates and temperatures
(Parodi, 1938; Knapp, 1990). The low temperature requirement evidenced by some lines
may thus constitute an advantage for materials adapted to saline environments. Future
studies on the germination behaviour of this species should involve populations from a
wide range of environmental conditions (i.e. contrasting origin associated with the soil
salinity) thereby allowing for the development of a broader appreciation of the 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.
References
Andrs, 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.
Beckstead, J., Meyer, S.E. and Allen, P.S. (1995). Effects of afterripening on cheatgrass (Bromus tectorum)
and squirreltail (Elymus elymoides) germination. Proceedings: Wildland Shrub and Arid Land Restoration
Symposium, (eds. B.A. Roundy, E.D. McArthur, J.S. Haley, and D.K. Mann), pp. 165-172, U.S. Department
of Agriculture, Forest Service, Ogden, UT.
Cceres, L. (1980). Caracterizacin climtica de la Provincia de Santa Fe. [Climatic characterization of the
Province of Santa Fe]. Santa Fe, Argentina, Direccin General de Suelos y Aguas. Ministerio de Agricultura
y Ganadera de la Provincia de Santa Fe, 21 pp.
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.

249

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

Espino, L.M., Seveso, M.A. and Sabatier, M.A. (1983). Mapa de suelos de la Provincia de Santa Fe. [Map of
soils of the Province of Santa Fe]. Rafaela, Argentina, Ministerio de Agricultura y Ganadera de la Provincia
de Santa Fe INTA.
Fandrich, L. and Mallory-Smith, C. (2005). Temperature effects on jointed goatgrass (Aegilops cylindrica) seed
germination. Weed Science, 53, 594-599.
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 center
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 y E. scabriglumis.
Seed Science and Technology, 25, 343-350.
Knapp, S.J. (1990). Recurrent mass selection for reduced seed dormancy in Cuphea laminuligera and Cuphea
lanceolata. Plant Breeding, 104, 46-52.
Maguire, J.D. (1962). Speed of germination aid in selection and evaluation for seedling emergence and vigor. Crop
Science, 2, 176-177.
Panigatti, J.L. (1968). Consideraciones sobre el clima del centro oeste de la Provincia de Santa Fe. [Considerations
about the climate in the west and center regions of the Province of Santa Fe]. Estacin Experimental
Agropecuaria Rafaela (INTA). Boletn Interno de Divulgacin, 15, 28 pp.
Parodi, L.R. (1938). El proceso biolgico de la domesticacin vegetal. [The biologic process of vegetal
domestication]. Revista Argentina de Agronoma, 5, 1-24.
Ranmuno, J.N. (1980). Forrajeras cultivadas para suelos salinos y/o alcalinos. [Forage species cultivated in
saline and/or alcaline soils]. Estacin Experimental Agropecuaria Marcos Jurez (INTA). Hoja informativa,
64, 5 pp.
Sen, D.N. and Mohammed, S. (1994). General aspects of salinity and the Biology of saline Plants. In: Handbook
of Plant and Crop Stress, (Ed. Mohammad Pessarakli), pp 125-148, New York.
Vargas Lpez, E. (1982). El agropiro criollo en la regin semirida pampeana. [The argentine wheatgrass in the
semiarid Pampas]. Estacin Experimental Agropecuaria Anguil (INTA). Informe de Tecnologa Agropecuaria
para la regin semirida pampeana, 78, 1-3.
Zabala, J.M., Tomas, P.A., Schrauf, G.A, and Giavedoni, J.A. (2009). Seed dormancy in Elymus scabrifolius
(Dll) J.H. Hunz. Seed Science and Technology, 37, 248-251.

250