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Journal of
Arid
Environments
Journal of Arid Environments 69 (2007) 169–176
www.elsevier.com/locate/jaridenv

Short communication

Seed germination of Trichocereus terscheckii

(Cactaceae): Light, temperature and
gibberellic acid effects
P. Ortega-Baesa,b,, M. Rojas-Aréchigab
a
Laboratorio de Investigaciones Botánicas (LABIBO), Facultad de Ciencias Naturales,
Universidad Nacional de Salta, Buenos Aires 177, 4400, Argentina
b
Departamento de Ecologı´a de la Biodiversidad, Instituto de Ecologı´a, Universidad Nacional Autónoma de México,
Apdo. Postal 70-275, 04510 México, D.F., México
Received 15 March 2006; received in revised form 22 July 2006; accepted 6 September 2006
Available online 3 November 2006

Abstract

In this paper, we evaluated the effect of light and temperature and addition of gibberellic acid
(GA3) in the germination of seeds of Trichocereus terscheckii in order to provide information about
germination requirements which could be use for conservation studies. The germination response
within a temperature gradient was evaluated for seeds arising from two populations: La Pedrera and
Cuesta del Obispo (Salta, Argentina). Seeds of T. terscheckii germinated within a range from 15 to
35 1C. Maximum germination percentages were found under white light and no germination was
recorded in darkness. GA3 at any concentration promoted germination either in white light or
darkness neither at a constant temperature nor at an alternating one.
r 2006 Elsevier Ltd. All rights reserved.

Keywords: Columnar cactus; Gibberellic acid; Photoblastism

1. Introduction

Diverse studies with Cactaceae have indicated that apart from humidity, light and
temperature are also important factors that can determine the germination of seeds

Corresponding author. Facultad de Ciencias Naturales, Universidad Nacional de Salta, Department of

Agronomy, Buenos Aires 177, 4400 Salta, Argentina. Tel.: +54 55 56229004.
E-mail address: portega@miranda.ecologia.unam.mx (P. Ortega-Baes).

0140-1963/$ - see front matter r 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jaridenv.2006.09.009
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170 P. Ortega-Baes, M. Rojas-Aréchiga / Journal of Arid Environments 69 (2007) 169–176

(Nobel, 1988; Rojas-Aréchiga and Vázquez-Yanes, 2000). With respect to light effect,
studies have demonstrated that barrel and globose cacti are positive photoblastic (Rojas-
Aréchiga et al., 1997; Bowers, 2000; Benı́tez-Rodrı́guez et al., 2004), while columnar cacti
may be indifferent to light or positive photoblastic (Rojas-Aréchiga et al., 1997, 2001;
Ramı́rez-Padilla and Valverde, 2005).
Several species from arid zones have shown higher germination percentages under
alternating temperatures than under constant temperatures (Mahmoud et al., 1983, 1984),
but for some cacti seeds it has been demonstrated that at a constant temperature between
20 and 30 1C, seeds reached their maximum germination (Rojas-Aréchiga et al., 1998;
Rojas-Aréchiga and Vázquez-Yanes, 2000). Hence, the use of alternating temperatures to
promote germination in cactus seeds has been demonstrated for some species (Fearn, 1981;
Rojas-Aréchiga et al., 1998; Yang et al., 2003), for other species, alternating temperatures
did not enhance or even reduced germination percentages compared to constant
temperatures (Rojas-Aréchiga and Vázquez-Yanes, 2000; Benı́tez-Rodrı́guez et al.,
2004). With respect to positive photoblastic seeds, the use of alternating temperatures
has not substituted for the light requirement for germination (Rojas-Aréchiga et al., 2001;
Benı́tez-Rodrı́guez et al., 2004).
The effect of gibberellic acid (GA3) on promoting germination of seeds of Cactaceae
has been poorly studied and results are diverse. Alcorn and Kurtz (1959) and Mc Donough
(1964), asserted that GA3 at 500 and 1000 ppm promoted germination of seeds under
white light and darkness. On the other hand, for Melocactus caesius, Stenocereus stellatus
and Mammillaria species, addition of gibberellic acid did not promote germination
in darkness (Arias and Lemus, 1984; Rojas-Aréchiga et al., 2001; Rojas-Aréchiga,
unpubl. res.).
Trichocereus terscheckii (Pfeiff.) Britton & Rose is a columnar cactus that grows in
the arid lands of Bolivia and Argentina (Kiesling, 1999). The establishment of individuals
is made by seed germination, hence no vegetative reproduction occurs. The region where
this species grows is threatened by loss and habitat degradation due to agricultural
expansion, wood extraction, and cattle overgrazing. Wood utilization of this species has
grown as a respond to an increasing tourism which demands the purchase of crafts
made with the wood obtained from this species. Species extraction impact has not been
assessed and at present no management program exists to control these activities in
northern Argentina. Seed germination studies are key tools in conservation programs
because they can be used for management programs and species reintroduction
(Ortega-Baes et al., 2005).
The aim of our study was to determine the effect of light, temperature and gibberellic
acid upon the seed germination of T. terscheckii. To fulfill this purpose, we formulated
the following questions: (1) Will T. terscheckii show either a positive photoblastic or
an indifferent behavior to light? (2) Will the use of an alternating temperature and the
addition of GA3 enhance germination in T. terscheckii seeds? (3) Will temperature
extremes affect seed germination? and (4) Will seeds arising from two populations
with different environmental traits show distinct temperature requirements for germina-
tion? Particularly for the germination response at different temperatures, we expect
T. terscheckii to be affected by temperature extremes. The maximum germination would be
obtained along a temperature gradient that varies among populations which grow within a
different temperature range. So, seeds that grow in areas with a wider thermal range,
would germinate within a wider temperature range.
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2. Materials and methods

2.1. Seed collection

Seeds were collected from 16 mature fruits from 10 individuals from La Pedrera site in
December 2000 (see Table 1). In the laboratory, seeds were extracted from the fruits and
put inside paper bags and kept in darkness at room temperature (2072 1C) until sowing
(February 2001). These seeds were used for the experiment in which the effect of light and
GA3 treatments were evaluated. In other experiment, we assessed the effect of temperature
on germination within a range from 10 to 40 1C every 5 1C. For this, seeds were collected in
December 2001 from 12 mature fruits obtained from 12 individuals from two sites: La
Pedrera and Cuesta del Obispo (see Table 1). For this experiment, seeds were sown in
January 2002.

2.2. Light treatments

To determine the effect of light quality upon the germination of seeds we made two
experiments consisting in a complete random design with five replicates per treatment
which were the following: white light, far-red light and darkness. For white light and far-
red light treatments we followed the methodology already used by Rojas-Aréchiga et al.
(1997). For darkness treatments, Petri dishes were wrapped with two aluminum foil sheets.
Each light treatment was exposed to a constant temperature of 25 1C and to an alternating
temperature 25/35 1C, with a thermoperiod of 16/8 h for the alternating temperature and a
photoperiod of 12 h for both temperature treatments. For each replicate, fifty seeds were
placed in Petri dishes with agar at 1% in distilled water and were put inside germination
chambers (Lab-Line Instruments, Inc., Model 844, Melrose Park, IL. USA).

2.3. Light and GA3 treatments

To determine the effect of gibberellic acid (GA3; Sigma G-7645 Gibberellin 90%) at
three concentrations (0, 500 and 1000 ppm) in white light and darkness we made two
experiments with a complete random design with four replicates. Each replicate consisted

Table 1
Geographic locations, altitude, mean annual precipitation, temperature, and vegetation type in La Pedrera and
Cuesta del Obispo (Salta, Argentina)

La Pedrera Cuesta del Obispo

Latitude 241 510 251 100

Longitude 651 210 651 490
Altitude (masl) 1729 2790
Mean annual precipitation (mm) 6407165 6677166
Mean annual temperature (1C) 16.8 10.2
Mean temperature in summer (1C) 21.9 14.6
Mean temperature in winter (1C) 10.5 4.4
Vegetation type Chaco forest Prepuna
Seed size (mg7S.E.) 0.7770.1 0.6270.15

Seed size is indicated (n ¼ 50). Climatological data obtained from Bianchi and Yañez (1992) and Bianchi (1996).
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of 50 seeds sown inside Petri dishes with agar at 1% in distilled water. The treatments were
the following: (1) white light without GA3, (2) white light with GA3 at 500 ppm; (3) white
light with GA3 at 1000 ppm; (4) darkness without GA3; (5) darkness with GA3 at 500 ppm,
and (6) darkness with GA3 at 1000 ppm. Seeds sown in Petri dishes were put inside
germination chambers at 25 1C and at an alternating temperature of 25/35 1C (thermo-
period 16/8 h), with a 12 h photoperiod.

2.4. Temperature treatments

With the aim to assess the temperature effect on the germination of seeds arising from
two different populations (see Table 1), we made a factorial experiment in which fifty seeds
were sown inside each of four Petri dishes on the surface of 1% agar in distilled water.
Dishes were incubated in growth chambers at seven temperatures ranging from 10 to
40 1C, every 5 1C (i.e. 10, 15, 20, 25, 30, 35 and 40 1C) with a 12 h photoperiod.
All experiments were followed for 30 days and germination was considered once the
radicle emerged from the testa. The response variables were the proportion of germinated
seeds at the end of each experiment.

2.5. Statistical analysis

For light and temperature treatments, data did not fulfill the basic assumptions to carry
out an ANOVA, so a Kruskal–Wallis test was carried out (Zar, 1996). In addition, for the
light experiments Mann–Whitney test were used to compare differences between
treatments in which germination occurred (Zar, 1996).
For the temperature experiment, a second analysis included treatments from 15 to 35 1C,
where germination data obtained as percentages were normalized by an arcsine square-
root transformation and were tested for statistical significance using an ANOVA (Zar,
1996). Means were compared by the LSD test at pp0.05 with the statistical program
Statistica software package (StatSoft, Inc., 1993, version 4.3).

3. Results

3.1. Light treatments

Seed germination was significantly different among light treatments at 25 1C (KW ¼

12.99; df ¼ 2; p ¼ 0.002) and at 25/35 1C (KW ¼ 13.06, df ¼ 2, p ¼ 0.001;
Fig. 1). Germination proportion was significantly different between white light and far-
red light treatments (25 1C: U ¼ 25, df ¼ 1, p ¼ 0.009; 25/35 1C: U ¼ 25, df ¼ 1,
p ¼ 0.008). Any seed germinated in darkness and maximum germination proportion
was obtained under white light at both constant and alternating temperature treatments
(Fig. 1).

3.2. Light and GA3 treatments

Seed germination showed significant differences among treatments at 25 1C (KW ¼

24.43, df ¼ 5, po0.001) and at 25/35 1C (KW ¼ 25.66, df ¼ 5, po0.001). Maximum
germination proportion was reached at white light with GA3-0 ppm at 25 1C, although no
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1
25 °C 25/35 °C

Germination (proportion)
0.9
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
0
White light Far-red light Darkness

Fig. 1. Proportion (mean7S.E.) of T. terscheckii seeds germinated under different light treatments at 25 and 25/
35 1C with a 12 h photoperiod.

Table 2
Proportion (mean7S.E.) of T. terscheckii seeds germinated under light and GA3 treatments

Treatments 25 1C 25/35 1C

Light regime GA3 (ppm)

White light 0 0.7670.029 0.5370.036

500 0.7370.034 0.5370.044
1000 0.7370.027 0.6670.028

Darkness 0 0 0
500 0 0
1000 0.00870.005 0.01670.012

significant differences were found among treatments in white light (KW ¼ 4.31, df ¼ 2,
p ¼ 0.12). For 25/35 1C in white light we found slightly significant differences among
treatments (KW ¼ 5.47, df ¼ 2, p ¼ 0.065), with the maximum germination proportion
with GA3-1000 ppm. Although the differences between temperature experiments could
not be proven statistically, the overall germination was higher at 25 1C than at 25/35 1C
(Table 2).
Although we did not find significant differences among treatments in darkness (25 1C:
KW ¼ 4.31, df ¼ 2, p ¼ 0.12; 25/35 1C: KW ¼ 4.29, df ¼ 2, p ¼ 0.12), a low germination
proportion was only achieved with GA3 at 1000 ppm (Table 2).

3.3. Temperature treatments

Seed germination was significantly different among treatments (KW ¼ 49.97, df ¼ 13,
po0.0001). Any seed germinated in 10 1C and 40 1C. For 15 to 35 1C treatments, seed
germination was significantly different between populations (F ¼ 16.99, df ¼ 1,
p ¼ 0.0003), among temperature treatments (F ¼ 26.33, df ¼ 4, po0.0001) and the
interaction was significant, too (F ¼ 2.88, df ¼ 4, p ¼ 0.04). Multiple range analysis did
not show significant differences between 15, 20 and 25 1C for both populations, but showed
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Cuesta del Obispo La Pedrera

Germination (proportion) 1

0.8

0.6

0.4

0.2

0
10 15 20 25 30 35 40
Temperature (°C)

Fig. 2. Germination proportion of T. terscheckii seeds from two populations within a temperature range from 10
to 40 1C.

significant differences at 30 1C and at 35 1C comparing with the other temperature

treatments, for both populations, too (Fig. 2).

4. Discussion

Results obtained with respect to light treatments, indicate that the seeds of Trichocereus
terscheckii showed a positive photoblastic response, similar to other columnar cacti
inhabiting arid and semiarid zones (Rojas-Aréchiga, et al., 1997; Rojas-Aréchiga and
Vázquez-Yanes, 2000; De la Barrera and Nobel, 2003). Although these seeds showed a
light requirement for germination, they are able to germinate under far-red light at lower
percentages than those obtained under white light, suggesting that they can germinate if
they come slightly buried in the soil where humidity may be available for a longer period of
time than in the soil surface. Light response of seeds can control the timing of germination
in their natural habitat, which may be crucial for seedling survival.
Bowers (2000) has considered that plant species which possess small and photoblastic
seeds are capable to form a seed bank. This author demonstrated a between-year seed bank
formation for Ferocactus wislizeni which possess small, positively photoblastic seeds. Also,
a short-term persistent bank has been demonstrated for T. atacamensis, a species with
small seeds and enforced dormancy, which also inhabits arid and semiarid zones at
northwestern Argentina (de Viana, 1999). Thus, the small, positively photoblastic seeds of
T. terscheckii should be capable of forming a seed bank; however this needs further field
investigation.
Seeds of T. terscheckii germinated to high percentages at temperatures between 15 and
30 1C, similar to those of several cacti species and did not germinate at the extreme
temperatures of 10 and 40 1C like other cacti species (Nobel, 1988; Rojas-Aréchiga et al.,
1998; Rojas-Aréchiga and Vázquez-Yanes, 2000). The results obtained in this study with
respect to temperature effects demonstrate that probably temperature does not limit the
distribution range of this species. Seeds from both populations showed a wide response to
temperature and the same behavior at the temperature extremes. However, assuming that
there may be other more important environmental factors regulating the germination of
this species.
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Alternating temperatures may (Fearn, 1974, 1981) or may not (Rojas-Aréchiga et al.,
1998; Ramı́rez-Padilla and Valverde, 2005) promote seed germination of cacti. For
T. terscheckii seeds, the alternating temperature used did not promote seed
germination neither in white light nor in darkness experiments compared to the constant
temperature. However, the use of another temperature regime concerning lower
temperatures (e.g. 10/20 1C), would throw different results.
We found no germination in darkness at the alternating temperature used suggesting
that this temperature did not substitute the light requirement for germination. These
results agree with those obtained for Stenocereus stellatus (Rojas-Aréchiga et al., 2001) and
with four Mammillaria species (Benı́tez-Rodrı́guez et al., 2004).
The addition of GA3 at two concentrations did not enhance seed germination of
T. terscheckii neither under white light nor under darkness conditions, which coincides
with the results obtained in other cacti species (Arias and Lemus, 1984; Rojas-Aréchiga et
al., 2001; Olvera-Carrillo et al., 2003). These contrasts with results shown by some authors
(Alcorn and Kurtz, 1959; Mc Donough, 1964; Deno, 1994). Lack of a response to GA3 and
the fact that fresh seeds were used in our study suggest that seeds of T. terscheckii do not
have any physiological dormancy. Thus, seeds are non dormant but have a light
requirement for germination and germinate over a wide range of temperatures. This
laboratory germination results can be applied to propagation projects that would support
conservation programs within the study site.

Acknowledgements

We are grateful to the anonymous referees for reviewing the manuscript. We thank
S. Sühring who helped us with the statistical analyses and A. Batis, M. Saravia, and
L. Hernández for technical assistance. POB thanks to Latin American Botanical Network
(RLB) for supporting his staying at UNAM, Mexico. MRA thanks the support given by
the Laboratorio de Dinámica de Poblaciones, Instituto de Ecologı́a, UNAM, México.

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