J.M. ZABALA, P. WIDENHORN AND J.F.

PENSIERO

Zabala, J.M., Widenhorn, P. and Pensiero, J.F. (2011), Seed Sci. & Technol., 39, 338-353

Germination patterns of species of the genus Trichloris in arid

and semiarid environments
J.M. ZABALAa, P. WIDENHORNa AND J.F. PENSIERO
Facultad de Ciencias Agrarias, Universidad Nacional del Litoral, Kreder 2805, Esperanza (S3080HOF)
Argentina. aex aequo (E-mail: jmzabala@fca.unl.edu.ar)

(Accepted March 2011)

Summary
Trichloris is an American genus with two species: T. crinita (Lag.) Parodi and T. pluriflora E. Fourn. Both
species are perennial warm season grasses and valued as forage. The objective of our study was to determine the
effect of different temperatures, seed weight and maternal effect (i.e., the contribution of the female parent to the
offspring phenotype) on the germination pattern in populations of Trichloris species from Argentina. The optimal
germination temperature for T. crinita ranged from 28 to 35C, whereas for T. pluriflora it was near 28C. In
all temperature treatments, the seeds of T. crinita populations showed higher values of germination than those
of T. pluriflora. Differences in seed weight were significant only among populations and there was a positive
correlation between germination and seed weight at suboptimal temperatures. These germination patterns may
explain the different distribution of species. There was no maternal effect on germination behaviour. Populations
with higher seed weight would be valuable if incorporated into breeding programmes.

Introduction
Two thirds of Argentina are characterised by arid and semi-arid rangeland ecosystems,
where, among other factors, deforestation and overgrazing have contributed to the
environmental and social-economic degradation process (Guevara et al., 1997). The
search for plants to be used in restoration projects and improve animal production is a
high priority in these ecosystems (Guevara et al., 2008).
Trichloris (Chloridoideae, Poaceae) is an American genus with two species: T. crinita
(Lag.) Parodi and T. pluriflora E. Fourn. Both species are perennial warm season grasses,
typical of the arid and semiarid environments of America, except for Canada (Peterson,
2001). Both are wide-spread in Argentina (Parodi, 1919; Zuloaga et al., 1994). Trichloris
crinita is found frequently in the arid, semiarid, sub-humid and humid environments
(Ragonese, 1956; Covas, 1978; Cavagnaro, 1988). On the other hand, Trichloris pluriflora
is found frequently in the semiarid, sub-humid and humid environments.
Trichloris crinita is an outstanding species due to its production, palatability and forage
quality in environments under drought stress (Bragadn, 1959; Wainstein and Gonzlez,
1971; Cerqueira et al., 2004). It has been selected as a priority species for restoration
programmes on degraded grasslands (Ragonese, 1956; Guevara et al., 2008). It is tolerant
to defoliation (Cavagnaro and Dalmasso, 1983), drought (Greco and Cavagnaro, 2003)
338

GERMINATION PATTERNS OF GENUS TRICHLORIS

and salinity (Aronson, 1989). Trichloris pluriflora, the species less studied, is abundant in
open forest lands, in places with a high availability of light (Morello and Saravia Toledo,
1959; Bordn, 1981). Some authors have emphasized its potential as forage (Daz, 1965;
Kunst et al., 1995).
Factors affecting the germination of seeds determine the success of programmes for
domestication of wild species (Casler and van Santen, 2010) and the sowing strategies for
restoration (Bischoff et al., 2006), and constitute an important aspect in the conservation
and regeneration of germplasm banks (Plucknett, 1992). Some of the most important
factors are: presence of seed dormancy, range of temperatures for optimal germination,
environmental maternal effects and seed weight.
Wild plant species have germination behaviour that varies among populations
(Stebbins, 1971; Meyer and Monsen, 1991; Zabala et al., 2009b). In many cases differences observed are attributed to different temperature requirements for germination or
seed dormancy (Simpson, 1990; Benech-Arnold et al., 2000).
The maternal effect is defined as the contribution of the female parent to the offspring
phenotype, assuming a similar chromosomal contribution between parents (Roach and
Wulff, 1987). One of the characteristics mostly influenced by maternal environment is
germination behaviour (Simpson, 1990; Baskin and Baskin, 1998), although variation in
their effect has been found among species (Bischoff et al., 2006) and among populations
within species (Schmitt et al., 1992).
Seed weight is a key aspect in the germination behaviour and seedling growth in
wild plant populations (Harper, 1977; Westoby, 1998; Thompson et al., 2001; Wang et
al., 2009). The role of seed weight has been examined at the intra-population (Gross and
Smith, 1991; Milberg et al., 1996; Baloch et al., 2001), inter-population (Greipsson and
Davy, 1995; Qiu et al., 2010) and inter-specific levels (Thompsom, 1990; Milberg and
Lamont, 1997; Liu et al., 2007). In all cases, the relationship between seed weight and
germination behaviour is variable and depends on the level of analysis (i.e. species level,
population level, etc.), and the specific conditions of germination. Most plants show an
increase in seed weight through the domestication process and this contributes to a higher
seedling growth rate and more uniform germination (Doganlar et al., 2000). However,
many warm season forage grasses with small seeds still retain wild characteristics.
Because of this, seed weight is a primary selection criterion in forage breeding (Casler
and van Santen, 2010).
There is scarce information on the germination conditions for T. crinita and T.
pluriflora. Cabeza et al. (1999) recorded value of 98% of germination for a population
of T. crinita from Argentina under an alternating of temperature of 15-25C (15 and 9 h,
respectively). Marone et al. (2000) analyzed germination behaviour in a population of
the same species under a constant temperature of 30C and recorded germination value
of 99%. Pezzani and Montaa (2006) reported germination near 80% under alternating
temperature (27-33C; 10 and 14 h, respectively) for a population of T. crinita from
Mexico.
The objective of this study was to determine the effect of seed weight, maternal
effect on seeds and different temperatures on the germination pattern of T. crinita and T.
pluriflora populations from Argentina.
339

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

Materials and methods

Plant materials
Populations of T. crinita came from arid and semiarid environments, and populations
from T. pluriflora were from semiarid environments (table 1). Seeds of 12 populations,
six T. crinita and six T. pluriflora, were used in trial 1. Seeds of six populations, three
T. crinita: 7320, 7413 and 7602, and three T, pluriflora: 7460, 7483 and 7491 were used
in trial 2.
Germination Assays
Two germination trials were carried out. In trial 1, seeds listed in table 1 were collected
from their native habitat and were germinated at five constant temperature regimes, 15C,
20C, 28C, 35C and 42C in the dark. In trial 2, seed harvested from plants grown in
a common environment without limitations of water or nutrients (Esperanza, Santa Fe,
S 31 25'; W 60 56') were germinated at three constant temperature regimes, 15C, 28C,
42C and alternating temperatures 15-28C (12 h and 12 h, respectively). In the two trials,
germination tests were performed on freshly matured caryopsis (1 month before harvest).
In Trichloris species the seeds are mature when endosperm becomes vitreous. In trial 2
panicle harvest was made 35-40 days after flowering in order to obtain mature seeds.
Germination procedure
In both trials, germination tests were conducted by placing 50 caryopsis in a petri dish
containing three Whatman paper discs moistened with distilled water. Four petri dishes
(replicates) were used for each treatment. Petri dishes were arranged at random in each
germination chamber. The number of germinated seeds was counted daily for 30 days. Only
seeds showing a 2 mm long radicle were considered germinated. Viability of non-germinated
caryopsis was determined at the end of the experiment according to methodology proposed
by Cabeza et al. (1999).
In both trials, fresh seed weight was determined by weighing four replicates of 200
caryopsis from each population. Mean seed weight was expressed as 1000-caryopsis weight
(mg/1000 caryopsis).
Data analysis
The coefficient of germination (CG), also referred to as speed of germination, was calculated
using the Maguire Index (Maguire, 1962). The final germination percentage (FGP) of
viable caryopsis and CG were subjected to a nested analysis of variance to determine the
significance of species, populations within species, temperatures, and the interactions among
them on germination. The statistical effect of populations was considered random, thus it
was used as a source of error to analyze the effect of the species. Differences between mean
values were tested for significance using Tukey test (p 0.05). Prior to analysis, FGP values
were arcsin transformed (Zar, 1996); however, untransformed means are reported in tables
and figures. The relative importance of single factors in analysis of variance in relation to
the total variation was estimated through the components of variance (Quinn and Keough,
2002).
340

Chaquea
(Occidental)

Monte

Monte

Monte

Chaquea
(Occidental)

Chaquea
(Occidental)

Chaquea
(Occidental)

Cahaquea
(Occidental)

Chaquea
(Occidental)

Chaquea
(Chaco Serrano)

TC8009

TC7375

TC73201

TC7946

TP74831

TP7599

TP74911

TP74601

TP7931

TP8053

Semiarid - Dry forest (with Schinopsis lorentzii, Ruprechtia apetala,

Ziziphus mistol, Acacia aroma and Prosopis torquata)

Semiarid - Algarrobal (xerophytic forest dominated by

Prosopis flexuosa. with Larrea divaricada, Bulnesia retama and
Capparis atamisquea)

Semiarid - Dry forest (with Schiniopsis lorentzii, Aspidosmerma

quebracho-blanco, Ziziphus mistol, Porlieria microphylla and Larrea
divaricata).

Semiarid - Ruderal (roadside with Cenchrus sp.)

Semiarid - Dry forest (with Schinopsis lorentzii, Prosopis sp.,

Ziziphus mistol, Cercidium praecox and Opuntia quimilo)

Semiarid - Dry forest (with Ruprechtia apetala, Prosopis torquata,

Bulnesia foliosa, Ximenia americana and Larrea sp.).

Arid - Jarillar (shrubland dominated by Larrea sp.)

Arid - Shrubland with Lycium sp., Schinus sp. and Larrea sp.

Arid - Saline shrubland with Atriplex sp., Allenrolfea vaginata and

Prosopis strombulifera

Semiarid - Algarrobal
(xerophytic forest dominated by Prosopis flexuosa.)

Semiarid - Quebrachal (xerophitic forest dominated by Aspidosperma

quebracho-blanco with Larrea divaricata and Larrea cuneifolia)

Semiarid - Salinas Grandes

(saline environment with halophytic vegetation)

Environment - Habitat

Populations used in trials 1 and 2; 2 According to Cabrera (1994); 3 Mean annual precipitations;
According to Instituto Nacional de Tecnologa agropecuaria INTA (http://geointa.inta.gov.ar/)

Chaquea
(Occidental)

TC74131

Chaquea
(Occidental)

Phytogeographic
province
(District)2

TC76021

ID

30 59' 12''

33 07' 08''

31 32' 7,6''

29 41' 43''

29 06' 23''

28 45' 35''

32 20' 02''

34 46' 19''

32 25' 10''

32 15' 20''

31 20' 38''

29 51' 00''

Lat.

Table 1. List of populations of Trichloris crinita and T. pluriflora and environmental conditions at the collection site.

65 04' 45''

66 31' 00''

67 16' 49''

66 56' 11''

36 05' 16''

66 01' 03''

67 17' 12''

68 36' 42''

68 36' 38''

65 09' 31''

66 36' 35''

64 40' 11''

Long.

684

573

442

477

579

471

333

329

150

667

471

683

Prec3
(mm)

Soil depth
(20 cm)

Climate

Climate

Climate

Climate

Climate

Climate and
soil salinity

Climatic

Climate and
soil salinity

Climate

Climate

Soil salinity

Principal
limitation to
plant growth4

GERMINATION PATTERNS OF GENUS TRICHLORIS

341

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

Seed weight was subjected to a nested analysis of variance to determine the significance
of species and populations within species. Association between seed weight and FGP and
CG were calculated using Pearsons correlation (p 0.05). Data were examined using
INFOSTAT statistical package (INFOSTAT version 2009).

Results
Trial 1 - Native Habitat
Variations in FGP and CG were highly significant (p < 0.001) for species, populations
within species, temperature and their interactions (table 2).The significant interactions
indicated that the caryopsis from different species and populations varied in germination
responses depending on the temperature treatment applied. Because of these significant
interactions it was concluded that temperature treatments had to be analyzed separately.
Table 2. Analysis of variance of final percentage of germination (FPG) and coefficient of germination (CG) in
trial 1.
DF1

Source of variation

FPG

CG

MS2

MS2

Species

23807.8

447.6*

519712.2

728.4*

Populations within species

2918.7

54.9*

73831.2

103.5*

Temperature

25396.5

477.5*

423487.1

593.6*

Species temperature

2092.3

39.3*

44953.3

63.0*

40

612.7

11.5*

7736.4

10.8*

Within populations (error)

180

53.2

Total

239

Populations within species Temperature

713.4

Degrees of freedom; Mean squares; * P < 0.001

There were significant differences among species, and populations within species
(p < 0.001) at all the temperatures tested (table 3). FGP and CG were higher in T. crinita
than in T. pluriflora in all temperature regimes (figures 1 and 2).
At 15C and 20C, the variation was mostly explained at the population level,
while at 28C, 35C and 42C the species component of variance was higher than that
from populations components (table 3). At 15 and 20C analysis of variance yielded
negative components of variance at species level. These components were replaced by
zero (Quinn and Keough, 2002). At 15C, the highest FGP for T. crinita was 51% in
population TC8009, with two populations having FGPs between 20 and 30%. However
three populations of T. crinita (TC7375, TC7413 and TC7602) and four populations of T.
pluriflora (TP7491, TP7460, TP7483 and TP7599) had an FGP lower than 3% (figure 1a).
At 20C, two populations of T. crinita (TC7320 and TC8009) had an FGP higher than
342

10
36

10
36

10
36

10
36

10
36

DF1

445.51
86.29

9544.8

674.1
54.2

17654.8

501.7
48.1

2001.7

2651.6
57.8

2639.3

1096.8
19.6

336.2

MS2

89.8
86.3

6066.2

154.9
54.2

11320.5

113.4
48.1

999.9

648.46
57.77

-8.19

269.3
19.6

-507.0

CV3

1.4
1.4

97.2

1.3
0.5

98.2

9.8
4.1

86.1

91.8
8.2

93.2
6.8

Percentage of
variation

Degrees of freedom; 2 Mean square; 3 Component of variance; * P < 0.001

Populations within sp.

Within pop. (error)

at 42C
Species

Populations within sp.

Within pop. (error)

at 35C
Species

Populations within sp.

Within pop. (error)

at 28C
Species

Populations within sp.

Within pop. (error)

at 20C
Species

Populations within sp.

Within pop. (error)

at 15C
Species

Source of variation

FPG

5.2*

110.6*

12.4*

325.6*

10.4*

41.6*

45.9*

45.7*

56.0*

17.11*

25161.4
1284.62

188876.2

20471.4
1194.4

334460.9

27017.4
683.9

138710.5

29487.8
325.7

36143.4

2638.8
78.6

1334.2

MS2

5969.2
1284.6

109143.2

4819.2
1194.4

209326.3

6583.4
683.9

74462.1

7290.5
325.7

4437.1

640.0
78.6

-869.7

CV3

CG

5.1
1.1

93.8

2.2
0.6

97.2

8.1
0.8

91.1

60.5
2.7

36.8

89.1
10.9

Percentage of
variation

19.6*

147.0*

17.1*

280.1*

39.5*

202.8*

90.5*

111.0*

33.5*

16.9*

Table 3. Analysis of variance and components of variance (CV) of final percentage of germination (FPG) and coefficient of germination (CG) in trial 1 (Native
Habitat) at different temperatures.

GERMINATION PATTERNS OF GENUS TRICHLORIS

343

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

85%, while two populations of T. crinita (TC7375 and TC7602) and three populations of
T. pluriflora (TP7483, TP7491 and TP7599) did not show FGP values higher than 10%
(figure 1b).
The highest values of FGP (figure 1c and d) and CG (figure 2c and d) were recorded
at 28C and 35C for T. crinita, and at 28C for T. pluriflora. In these optimal temperature
conditions, seed started to germinate 1-2 days after sowing and showed the maximum
FGP after 21 days in all populations (figure 3).

FPG (%)

a)

100
90
80
70
60
50
40
30
20
10
0

c) 100

LSD sp: 2.6 LSD pop: 10.9

LSD sp: 4.45 LSD pop: 18.8

LSD sp: 14.1 LSD pop: 17.1

LSD sp: 4.3

LSD pop: 18.2

d)

TC
7

3
TC 20
73
TC 75
74
TC 13
76
TC 02
79
TC 46
80
TP 09
74
TP 60
74
TP 83
74
TP 91
75
TP 99
79
TP 31
80
53

90
80
70
60
50
40
30
20
10
0

FPG (%)

b)

LSD sp: 5.4 LSD pop: 22.9

100
90
80
70
60
50
40
30
20
10
0

Populations

TC

73
TC 20
73
TC 75
74
TC 13
76
TC 02
79
TC 46
80
TP 09
74
TP 60
74
TP 83
74
TP 91
75
TP 99
79
TP 31
80
53

FPG (%)

e)

Populations
Figure 1. Mean ( S.D.) final germination percentages (FGP) for seeds of populations of T. crinita (black
columns) and T. pluriflora (grey columns) grown under different temperatures in trial 1. a) 15C; b) 20C;
c) 28C; d) 35C and e) 42C. Dashes lines indicate species means. LSD sp: Least significant difference between
species at p < 0.05, (Tukeys test). LSD pop: Least significant difference between populations.

344

GERMINATION PATTERNS OF GENUS TRICHLORIS

a)

400

LSD sp: 5.2 LSD pop: 21.9

LSD sp: 10.5 LSD pop: 21.9

b)

LSD sp: 15.3 LSD pop: 64.6

LSD sp: 20.2 LSD pop: 85.4

d)

350

CG

300
250
200
150
100
50
0

c)

400
350

CG

300
250
200
150
100
50

TC

e)

73
TC 20
73
TC 75
74
TC 13
76
TC 02
79
TC 46
80
TP 09
74
TP 60
74
TP 83
74
TP 91
75
TP 99
79
TP 31
80
53

400

LSD sp: 20.9 LSD pop: 88.6

350

Populations

CG

300
250
200
150
100
50
TC
73
TC 20
73
TC 75
74
TC 13
76
TC 02
79
TC 46
80
TP 09
74
TP 60
74
TP 83
74
TP 91
75
TP 99
79
TP 31
80
53

Populations
Figure 2. Mean (S.D.) coefficient of germination for seeds of populations of T. crinita (black columns) and
T. pluriflora (grey columns) exposed to different temperature treatments in trial 1. a) 15C; b 20C; c) 28C;
d) 35C and e) 42C. Dashed lines indicate species means. LSD sp: least significant difference between species
at p < 0.05, (Tukeys test). LSD pop: least significant difference between populations.

Seed weight differed significantly among populations within species (p < 0.001) (table
4), but it was not so among species (p = 0.68) (figure 4). Correlations coefficient were
significant (p 0.05) and positive between seed weight and both FGP and CG only at
15C and 20C (figure 5).
345

FPG (%)

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

TC7320 TC7413
TC8009
TC7375
TC7946
TC7602

100
90
80
70
60
50
40
30
20
10
0

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

TP7460
TP7483
TP7599
TP7491 TP7931 TP8053

0 2 4 6 8 10 12 14 16 18 20 22 24 26 28 30

Days
Days
Figure 3. Germination response curves for seeds of populations of T. crinita (left) and T. pluriflora (right) under
optimal temperature (28C) in trial 1 (Native Habitat). FPG: final percentage of germination.

Table 4. Analysis of variance of seed weight in trial 1 (Native Habitat).

DF1

MS2

0.1600

Populations within species

10

0.8900

Within populations (error)

36

0.0081

Total

47

Source of variation
Species

F
0.18 NS
110.16*

Degrees of freedom; 2 Mean squares; NS: not significant; * P < 0.001

Seed weight
(mg/1000 seeds)

200

LSD sp: 60.8 LSD pop: 22.3

175
150
125
100
75

TC
73
TC 20
73
TC 75
74
TC 13
76
TC 02
79
TC 46
80
TP 09
74
TP 60
74
TP 83
74
TP 91
75
TP 99
79
TP 31
80
53

50

Populations
Figure 4. Mean ( S.D.) seed weight of populations used in trial 1. Dashed lines indicate species means. LSD
sp: least significant difference between species at p < 0.05, (Tukeys test). LSD pop: least significant difference
between populations.

Trial 2 Common Environment

Germination patterns in trial 2 were similar to those in trial 1. In all temperature regimes,
the seeds of T. crinita populations showed higher values of FGP and CG than those
of T. pluriflora (figure 6). Differences were minimal following germination at 28C, in
346

GERMINATION PATTERNS OF GENUS TRICHLORIS

FPG (%)

a)

100

b)

r = 0.90

r = 0.89

TC 7320

TC8009

80
TC8009

60
TP 7599
TP 7491 TC 7602
TP 7483 TC 7413
TP 7460 TC 7375

40
20

TC 7320

TP 7931

TP 7599 TC 7602
TP 7491 TC 7413
TP 7483 TC 7375
TP 7460

TP 7931
TP 8053

TP 8053

TC 7946

TC 7946

c)

d)

250

CG

200
150
100
50

TC8009

r = 0.90

TC 7320

r = 0.86

TP 7931
TP 8053
TP 7599 TC 7602
TP 7491
TC 7413
TP 7483
TC 7375
TP 7460

TC 7320

TP 7599 TC 7602
TC 7375
TP 7491
TC 7413
TP 7483
TP 7460

TC8009
TP 7931
TP 8053

TC 7946

TC 7946

0
50

75

100

125

150

175

Seed weight (mg/1000 seeds)

200

50

75

100

125

150

175

200

Seed weight (mg/1000 seeds)

Figure 5. Relationship between seed weight and both final germination percentage (FGP) and coefficient of
germination in trial 1. a) Mean final percentage of germination at 15C; b) Mean final percentage of germination
at 20C; d) Mean coefficient of germination at 15C y d) Mean coefficient of germination at 20C. r = Pearsons
correlation coefficient.

which the highest values of FGP and CG were recorded in both species (figure 6c-d). As
compared with the treatment at 28C, the alternating temperature showed a decrease of
FGP and CG in almost all the populations tested, although it did not affect the FGP in the
TC7320 population (figure 6e-f).
The seed weight from populations undergoing similar growing conditions was always
higher than that of the same population in their environment of origin (figure 4 and 7).
Differences in seed weight were significant among populations (p < 0.01) (figure 7). As
in trial 1, there was a positive correlation between seed weight and FGP and CG only at
15C (figure 8).
Under optimal germination conditions, five populations two of T. crinita (TC7602,
TC7946) and three of T. pluriflora (TP7491, TP7931, TP8053) which retained approximately 20% of seed dormancy were detected in trial 1.

Discussion
There was no maternal effect on germination in Trichloris species. Optimal germination
temperature for T. crinita ranged from 28C to 35C, whereas for T. pluriflora it was
near 28C. Percentage germination at 28C would be recommended for germination tests
in both species. We showed that 7 and 21 days would be useful as the first and final
germination counts, respectively.
347

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

FPG (%)

e)

100
90
80
70
60
50
40
30
20
10
0

100
90
80
70
60
50
40
30
20
10
0

b)

400
350

LSD sp: 5.3 LSD pop: 2.2

300

CG

250
200
150
100
50
0
LSD sp: 18.8 LSD pop: 18.1

LSD sp: 9.4 LSD pop: 26.1

d)

LSD sp: 22.1 LSD pop: 14.1

400
350
300
250
200
150
100
50
0

400

f)

LSD sp: 22.9 LSD pop: 55.3

350
300
250
200
150
100
50
0

LSD sp: 19.5 LSD pop: 22.3

i)

CG

FPG (%)

g) 100
90
80
70
60
50
40
30
20
10
0

LSD sp: 28.1 LSD pop: 17.1

CG

FPG (%)

c)

100
90
80
70
60
50
40
30
20
10
0

CG

FPG (%)

a)

TC7320 TC7413 TC7602 TP7460 TP7483 TP7491

400
350
300
250
200
150
100
50
0

LSD sp: 39.4 LSD pop: 49.8

TC7320 TC7602 TC7413 TP7460 TP7483 TP7491

Figure 6. Mean (S.D.) final germination percentage (FGP), on the left, and coefficient of germination (CG),
on the right, for seeds of populations of T. crinita (black columns) and T. pluriflora (grey columns) exposed to
different temperature treatments in trial 2. a-b) 15C; c-d) 28C; e-f) 15-28C (12 hours and 12 hour respectively)
and g-h) 42C. Dashed lines indicate species means. LSD sp: least significant difference between species at
p < 0.05, (Tukeys test). LSD pop: least significant difference between populations.

348

GERMINATION PATTERNS OF GENUS TRICHLORIS

400

Seed weight
(mg/1000 seeds)

LSD sp: 18.1 LSD pop: 133.2

300

200

100

0
TC7320 TC7602 TC7413 TP7460 TP7483 TP7491

Populations
Figure 7. Mean ( S.D.) seed weight of populations used in trial 2. Dotted lines indicate species means. LSD
sp: least significant difference between species at p < 0.05, (Tukeys test). LSD pop: least significant difference
between populations.

a)

b)
120

r = 0.95

80
60
40
20

r = 0.96

100
TC 7320
TC 7491
TC 7483

TC7602
TC7413
TP7460

CG

PFG (%)

100

80
60
40
20

0
150 175 200 225 250 275 300 325 350 375 400

Seed weight (mg/1000 seeds)

TC 7320
TP7460
TC7413
TC7602
TC 7483
TC 7491

0
150 175 200 225 250 275 300 325 350 375 400

Seed weight (mg/1000 seeds)

Figure 8. Dispersion graphics of seed weight versus final germination percentage (FGP) and coefficient of
germination (CG) in trial 1 at 15C; a) Mean FPG (final percentage of germination); b) Mean coefficient of
germination. r = Pearsons correlation coefficient.

In general, populations of T. crinita germinated in a wider temperature range than

those for T. pluriflora. This could partly explain a wider geographical distribution
shown by T. crinita compared to that of T. pluriflora, although there is a great interpopulation variability, particularly under suboptimal temperature conditions. Germination
capacity is an indicator of the distribution and abundance of plants, though its relative
importance has been debated (Grime et al., 1981; Burke and Grime, 1996; Thompson et
al., 2001; Ramrez-Padilla and Valverde, 2005). Most work indicates that a wide range of
germination temperatures enhances seedling competitiveness; however in the long term,
other characteristics are also considered of a great importance.
The decrease of germination in suboptimal temperatures could be an ecological
adaptation to germinate under favourable conditions of soil moisture. Under the
environmental conditions of Chaquea and Monte Phytogeographic Provinces, soil
moisture is restrictive in winter for plant growth. Rains begin in spring (October) and
continue in summer until autumn (March-April).
349

J.M. ZABALA, P. WIDENHORN AND J.F. PENSIERO

Many wild species require specific germination conditions such as alternating

temperature (Baskin and Baskin, 1998; Zabala et al., 2009a). In spite of previous reports
for T. crinita (Cabeza et al., 1999; Pezzani and Montaa, 2006), our results showed that
there is no such requirement in either species of Trichloris.
The populations of T. crinita coming from saline environments (TC7602, TC7375
and TC7946) showed a narrow range of germination temperatures (figure 1 and 6), being
adaptive characteristics in halophyte species (Flowers et al., 1986). Even though the
germination behaviour of populations coming from a saline environment requires further
study, it seems that high temperature requirements for germination, which coincide with
the rainy season (i.e. decrease of soil salinity), would be a strategy of survival in that
environment.
Under suboptimal temperature conditions, the germination of T. crinita and T. pluriflora
was directly associated with seed weight. Increased seed weight was positively correlated
with cold temperature germination in Arabidopsis thaliana (Boyd et al., 2007), Festuca
halli (Qiu et al., 2010) and in other 131 plants species (Thompson, 1990). Other reports
indicated that in certain species seed weight is positively correlated with germination
capacity under other stressful conditions such as drought (Gross and Smith, 1991; Daws
et al., 2008) or nutrient deficiency in soils (Milberg and Lamont, 1997). These results
may be related to a greater capacity of reserve mobilization in bigger seeds under these
conditions (Kidson and Westoby, 2000), especially within the grass family which have
low embryo/reserve ratio (Martin, 1946). In general, when germination conditions are
optimal, the correlation between seed weight and germination behaviour is not so evident
(Wang et al., 2009) or shows opposite results (Susko and Lovett-Doust, 2000). Greipsson
and Davy (1995) studied germination behaviour under optimal temperatures in 34
populations of Leymus arenarius with a variation of seed weight by a factor of about ten.
They found a positive correlation between seed weight and final germination percentage,
but a negative correlation with regard to germination speed. Our results do not agree with
their findings, since no relation was found between seed weight and germination speed or
final germination percentage under optimal temperature regimes.
Populations with high seed weight were recorded in two species, three from T. crinita
and two from T. pluriflora, and come from different regions. This means that in the
populations analyzed, there is no consistent pattern between the seed weight and the
origin of the population, as has been demonstrated for other species (Baker, 1972).
Differences in seed weight were consistent both between populations harvested in their
original environment as well as those grown in a common environment. Regarding the
latter, seed weight was higher in all the populations and the range of differences was less.
The higher seed weight coming from plants growing in a common environment is possibly
due to the fact that seed production was performed under non-limiting environmental
conditions. These results suggest that both genetic and environmental factors affect the
weight of seed in T. crinita and T. pluriflora, as demonstrated in Andropogon hallii
(Glewen and Vogel, 1984), Panicum virgatum (Boe, 2003) and Elymus junceus (Berdahl
and Barker, 1984).
This study presented germination conditions for species of the genus Trichloris,
analyzing inter-specific and inter-population variability for the arid and semiarid regions
350

GERMINATION PATTERNS OF GENUS TRICHLORIS

of Argentina. Optimal germination temperatures were determined demonstrating the

presence of germination patterns that would partly explain the distribution of species in
Argentina. Inter-population differences in seed weight, associating greater weight with a
higher germination in suboptimal temperature regimes, were also identified. Populations
with higher seed weight would be valuable if incorporated into breeding programmes.

Acknowledgements
Financial support for this research was provided by Universidad Nacional del Litoral and
Agencia Nacional de Promocin Cientfica y Tecnolgica.

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