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Physiological dormancy and germination requirements of seeds of several

North American Rhus species (Anacardiaceae)
Xiaojie Li, Jerry M. Baskin and Carol C. Baskin
Seed Science Research / Volume 9 / Issue 03 / March 1999, pp 237 - 245
DOI: 10.1017/S0960258599000240, Published online: 22 February 2007

Link to this article: http://journals.cambridge.org/abstract_S0960258599000240

How to cite this article:
Xiaojie Li, Jerry M. Baskin and Carol C. Baskin (1999). Physiological dormancy and germination requirements of seeds of
several North American Rhus species (Anacardiaceae). Seed Science Research, 9, pp 237-245 doi:10.1017/
S0960258599000240
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Seed Science Research (1999) 9, 237245

237

Physiological dormancy and germination requirements of

seeds of several North American Rhus species
(Anacardiaceae)
Xiaojie Li,1 Jerry M. Baskin,*1 and Carol C. Baskin1,2
1
School of Biological Sciences, University of Kentucky, Lexington, Kentucky 40506-0225, USA; 2Department of
Agronomy, University of Kentucky, Lexington, Kentucky 40546-0091, USA

Abstract

Introduction

Fourteen seedlots of five species of Rhus were

surveyed for presence/absence of physiological
dormancy and/or for germination requirements of nondormant seeds. Physiological dormancy was present in
the four seedlots of R. aromatica studied, but not in
either of the two seedlots of its close relative R.
trilobata, which is in contrast to previous reports.
Neither were seeds of R. glabra, R. typhina, nor R.
virens physiologically dormant. Stratification at 5oC for
1 week or incubation in 500 or 1000 mg/l solutions of
gibberellic acid broke physiological dormancy in > 90%
of the R. aromatica seeds. Maturation desiccation acted
as a switch from a developmental to a germinative
mode in R. aromatica embryos, whereas it was not
required for germination of R. glabra or R. virens (R.
trilobata and R. typhina not tested). Seeds of all five
species incubated on a moist substrate became fully
imbibed in 2 d, at which time moisture content was
approx. 7080% of their initial weight. In general,
germination of non-dormant seeds was rather
insensitive to temperature and light. Seeds germinated
equally well in light and in darkness over a daily
(12 h/12 h) temperature range of 15/635/20oC. Over a
4 week period, the best germination percentages were
obtained at 25/15 and 20/10oC, whereas 35/20oC
appeared to be supraoptimal, though not always
significantly so. If the incubation period was extended
to 30 weeks, germination percentages were as high at
15/6oC as at 25/15 and 20/10oC.

Seeds of Rhus L. species (Anacardiaceae) have physical

(hard fruit coat) dormancy caused by a waterimpermeable endocarp, and some have physiological
dormancy as well. Much of the information on seed
germination in Rhus is on breaking endocarp
impermeability (Boyd, 1943; Lovell, 1964; Smith, 1970;
Heit, 1967; Farmer et al., 1982; Keeley, 1987, 1991). The
literature is confusing with regards to embryo
dormancy and to the germination requirements of
non-dormant seeds. For example, Norton (1985, 1986,
1987) demonstrated that seeds of R. typhina are
physiologically dormant, in stark disagreement with
reports by other investigators (e.g., Heit, 1967;
Brinkman, 1974; Li and Xu, 1989). Similarly,
stratification of scarified seeds was recommended for
R. virens by Tipton (1992), whereas Nokes (1986) stated
that scarified seeds of this species germinate without
being stratified. One of the factors contributing to such
conflicting reports is the possibility that embryo
dormancy could vary among seedlots (Heit, 1970).
Unfortunately, very few concrete data are available on
this subject. Germination of R. glabra and R. copallina
seeds was fairly insensitive to either light or
temperature (Lovell, 1964; Farmer et al., 1982).
However, Keeley (1987, 1991) reported that light
significantly increased germination percentage of R.
trilobata seeds.
Maturation desiccation is the terminal event of
seed development in many species. It was shown for
many cereals and legumes that the seed switches from
a developmental to a germinative mode during
maturation drying (Wellington, 1956; King, 1976;
Adams and Rinne, 1981; Long et al., 1981; Dasgupta
and Bewley, 1982; Bewley and Black, 1994).
Desiccation
results
in
the
cessation
of
developmentally-related synthetic events and the
onset of synthesis associated with germination and
postgerminative growth (Kermode and Bewley, 1985,
1989; Cornford et al., 1986; Miles et al., 1988; Bewley
and Oliver, 1992). However, no information is

Keywords: light and seed germination, imbibition,

physiological seed dormancy, Rhus spp., temperature and
seed germination

*Correspondence
Fax: 606-257-1717
Email: jmbask0@ukcc.uky.edu

238

X. Li et al.

available on Rhus with respect to the effects of

maturation desiccation on germination.
Thus, the objectives of the present study were to:
(1) Survey for presence/absence of physiological
dormancy in five North American Rhus species
and investigate methods to break it in seeds of
those species (if any) in which it is present.
(2) Test the effects of maturation desiccation on
subsequent germination.
(3) Determine the time course of imbibition prior to
germination, once physical dormancy is broken.
(4) Test the effects of temperature and light regimes
on germination of non-dormant seeds.

Materials and methods

Species and study system

The five species included in this study, R. aromatica, R.

glabra, R. trilobata, R. typhina, and R. virens, are shrubs
native to the United States and/or adjacent
Canada/Mexico. Rhus glabra occurs in all 48
contiguous states (Little, 1977), north to southern
Canada and south to northern Mexico (Barkley, 1937).
In contrast, the other four species have relatively
limited distributions. Rhus aromatica occurs in Quebec
and Ontario, Canada, south to Florida and west to
Kansas, Oklahoma, and Texas (Barkley, 1937). A
western USA relative of R. aromatica, R. trilobata (= R.
aromatica var. trilobata) is distributed from Alberta and
Saskatchewan, Canada, to Mexico (Barkley, 1937).
Rhus typhina ranges from Nova Scotia, Canada, to
Minnesota and Iowa and south to Texas, Louisiana,
and Florida (Barkley, 1937; Hall, 1977), and Rhus virens
from Texas and New Mexico, south to Mexico
(Barkley, 1937; Vines, 1960). All species are winterdeciduous except R. virens, which is evergreen.
Fruits of Rhus are 1-seeded drupes consisting of a
fleshy exocarp and mesocarp and a stony endocarp,
which encloses the anatomically-true seed. The
endocarp plus seed together act as the germination
unit. Thus, it is referred to as seed throughout this
paper. Seeds of R. glabra and R. typhina are approx.
3 mm by 2 mm with a mass of about 7 mg and are
much smaller and weigh less than those of R.
aromatica, R. trilobata, and R. virens, which are approx.
5 mm by 4 mm and have a mass of 1423 mg. The
endocarps of all five species are impermeable to water,
and thus have physical dormancy (Li et al., 1999a).
Seed collection and cleaning

To determine how widespread embryo dormancy is

among the five species, and among seedlots within R.
aromatica, R. glabra, and R. trilobata, mature fruits were
collected from two or more geographical locations

within their natural range; only one seedlot each was

available for R. typhina and R. virens. Only one seedlot
per species was used in studies on the requirements
for breaking physiological dormancy of R. aromatica
and on the requirements for germination of nondormant seeds of all five species. Details about
locations and dates of fruit collection as well as seed
cleaning procedures are described elsewhere (Li et al.,
1999a).
Germination procedures

Germination tests were performed in temperatureand light-controlled incubators at a 14 h daily

photoperiod (approx. 40 mol m2 s1, 400700 nm of
cool white fluorescent light). Seeds were incubated in
Petri dishes (1.5 cm 3 9 cm) on moist white quartz
sand. Unless otherwise specified, the dishes were
incubated at an alternating temperature regime
(12 h/12 h) of 25/15oC. The photoperiod began 1 h
before the beginning of the high temperature phase of
the thermoperiod and ended 1 h after the beginning of
the low temperature phase. Each germination
treatment or treatment-combination had four
replicates of 50 successfully-scarified (Li et al., 1999a)
seeds per dish. Unless otherwise specified, water was
added when needed to keep the sand in the Petri
dishes moist. To reduce water loss caused by
evaporation, the dishes were wrapped with plastic
film. Germinated seeds (i.e., those with radicles 2
mm in length) were recorded and removed from the
dishes weekly for a period of 4 weeks, except when
specified otherwise. Seeds that did not germinate were
identified as either imbibed and empty, imbibed and
filled, or non-imbibed (i.e., hard) at the end of each
germination test. Germination percentage (GP) was
based on imbibed and filled seeds only. We believe
that such GP values best indicate the extent to which
viable seeds potentially can germinate once rendered
permeable.
Detecting physiological dormancy

To determine if embryos of scarified seeds were

physiologically dormant or not, seeds were either
stratified (treatment) or not (control) prior to
incubation at 25/15oC in light. Seeds were cold
stratified on moist sand in Petri dishes at 5oC for 21
days, after which they were incubated at 25/15oC.
Effects of maturation desiccation on germination

To determine if maturation desiccation acts as a switch

from a developmental to a germinative mode in Rhus,
seeds of R. aromatica, R. glabra, and R. virens with
potentially germinable embryos (i.e., excised embryo
can germinate) were collected from the field prior to

Germination of Rhus seeds

maturation desiccation. Such seeds were divided into

two sets. One set of seeds was prepared immediately
for germination tests (not desiccated), while the other
set was allowed to dry under ambient laboratory
conditions for 7 days to simulate maturation
desiccation. After 7 days, the seeds became waterimpermeable and thus were physically dormant. They
were scarified and incubated as described previously.
As seeds of R. aromatica were shown to have
embryo dormancy (see Results), half of them were
stratified at 5oC for 21 days before placement in the
incubator for germination tests, while the other half
were not.
Effects of gibberellic acid (GA3) and stratification on
R. aromatica seeds

As R. aromatica is the only species in which embryo

dormancy was detected (see Results), subsequent
experiments in search of effective methods to break
this physiological dormancy were carried out for a
representative seedlot of this species only. H2SO4scarified seeds of seedlot Fky94 were treated as
follows: non-treated (control); stratified at 5oC for up
to 25 days; and soaked in GA3 (as 75% KGA3)
solutions prior to incubation. For GA3 treatments,
seeds were soaked in GA3 (United States Biochemical
Corporation, Cleveland, Ohio, USA) solutions of 2.5,
12.5, 25, 500, and 1000 mg/l for 24 h, placed on sand
moisten with the corresponding GA3 solutions, and
incubated. Dishes were wrapped with plastic film to
retard evaporation.
Water uptake during imbibition

To determine how much and how fast water is

absorbed by seeds during imbibition, water uptake of
25 scarified seeds each of R. aromatica, R. glabra, R.
trilobata, and R. virens was measured individually at
4-h intervals for the first 24 h and at daily intervals for
the following 5 days. Each individual seed was
weighed and then placed in a Petri dish on moist sand.
After each interval, the seed was taken out of the Petri
dish, quickly surface-dried with Kimwipe tissue
paper, weighed to the nearest 0.001 mg, and returned
to the dish. Seeds with radicles 2 mm were
considered to be germinated and thus discontinued
for the imbibition tests. Percentage water uptake was
calculated as amount of water taken up relative to
original weight of the seed.
Effects of light and temperature on germination of
non-dormant seeds

In these experiments, germination tests were

conducted in either white light or in constant darkness
at daily temperature regimes of 35/20, 30/15, 25/15,

239

20/10, and 15/6oC. These temperatures approximate

the mean monthly maximum and minimum air
temperatures in Kentucky and Tennessee during the
growing season (Wallis, 1977): July and August, 35/20;
June and September, 30/15; May, 25/15; April and
October, 20/10; and March and November, 15/6.
Darkness was provided by wrapping the Petri dishes
with two layers of aluminium foil. All germination
tests were run for a period of 4 weeks, except at
15/6oC, where seeds were incubated for 30 weeks, by
which time all viable seeds had germinated.
The effect of red light and of far-red light on
germination of R. glabra seeds also was investigated at
30/15oC by exposing them to 14 h red- or far-red light
and 10 h of darkness per day for 4 weeks. The red- and
far-red light sources were phosphor type 236 and
232 fluorescent tubes, respectively (GTE Products
Corporation, Danvers, Massachusetts, USA). The red
light produced energy at 600-730 nm, with peak
emission at 660 nm; photon flux density at seed level
was approx. 27 mol m2 s1. The ratio of far-red
(700800 nm) to red (600700) photon irradiance was
approx. 0.2. The far-red light produced energy at
600940 nm, with peak emission at 740 nm; photon
flux density at seed level was approx. 15 mol m2 s1.
The ratio of far-red (700800 nm) to red (600700 nm)
photon irradiance was approx. 8.7.
Data analysis

All germination percentage data were rounded to

the nearest whole number, and water uptake data
were rounded to one decimal place. Data were
analyzed by either a one-way or a two-way ANOVA
followed by Tukeys multiple comparison test at
P 0.05 (SAS Institute, 1989). Square roots of all
percentage data were arcsine-transformed prior to
statistical analyses.

Results
Seeds of all four seedlots of R. aromatica were
physiologically
dormant;
non-stratified
seeds
germinated from 0 to approx. 15% and stratified seeds
to >90% (Table 1). In contrast, non-stratified seeds of
R. glabra germinated from 74% to 93%, and these
percentages were statistically equivalent to those
obtained for stratified seeds. Germination percentages
of stratified and of non-stratified seeds of R. trilobata,
R. typhina, and R. virens also did not differ.
Non-stratified seeds of R. aromatica with a
potentially germinable embryo, collected prior to
maturation desiccation (i.e., before they became waterimpermeable), germinated to only 2 1%, whereas
stratified seeds germinated to 25 9% (Table 2).
However, if seeds first were surface-dried in the

240

X. Li et al.
Table 1. Effect of cold stratification on germination of scarified seeds1 of five Rhus
species. For each seedlot, data within a row with the same superscript do not differ
statistically (Tukeys test, P 0.05).
% germination after 4 weeks of
incubation at 25/15oC in light
Species

Seedlot2

Non-stratified

Fky94
Mtn96
Rtn89
Rva96
Cky94
Fky93
Fky94
Hky94
Rky93
Wtn94
Snb963
Sut94
Ctn96
Btx96

16 1a
7 2a
7 4a
0 0a
74 3a
93 1a
82 3a
81 3a
92 2a
84 3a
54 8a
93 2a
99 1a
92 3a

21-day stratified

Rhus aromatica

R. glabra

R. trilobata
R. typhina
R. virens

95 2b
98 1b
91 2b
94 5b
78 2a
92 1a
88 1a
82 2a
96 1a
88 3a
65 14a
95 1a
99 1a
94 2a

Seeds of R. aromatica, R. virens, and R. trilobata were scarified by soaking them in

concentrated H2SO4 for 1.0, 1.0, and 0.5 h, respectively, and those of R. glabra and
R. typhina were scarified by immersing them in boiling water for 1 min (Li et al., 1999a).
2
See Li et al. (1999a) for details of locations and collection dates.
3
Many seeds were heavily-infested with mold, thus accounting for the moderate
germination percentages.

Table 2. Effect of maturation desiccation on subsequent germination of three Rhus

species. For each species, data within a row with the same superscript do not differ
statistically (Tukeys test, P 0.05).
% germination of freshly-collected, permeable seeds1
after 4 weeks of incubation at 25/15oC in light
Non-desiccated

Desiccated2

Rhus aromatica:
Non-stratified
Stratified

2 1a
25 9a

20 5b
76 4b

R. glabra:
Non-stratified

89 11a

86 8a

R. virens:
Non-stratified

97 2a

95 4a

Species

Immature seeds were collected before the endocarp became impermeable, but after
excised embryos were capable of germinating.
2
Freshly-collected, permeable seeds were allowed to surface-dry in the laboratory for
7 days, until they became water-impermeable (to simulate the maturation desiccation
process in the field). Hard seeds then were scarified by soaking those of R. aromatica and
of R. virens in concentrated H2SO4 for 1 h and those of R. glabra in boiling water for 1 min,
prior to stratification treatments/germination tests.

laboratory until they became impermeable to water,

germination percentages of non-stratified seeds
increased significantly to 20 5% and those of
stratified seeds to 76 4%. On the contrary, freshly-

collected, permeable seeds of both R. glabra and

R. virens germinated to 89 11% and 97 2%,
respectively, without stratification, which was not
significantly different from germination percentages of

Germination of Rhus seeds

Table 3. Effects of gibberellic acid (GA3) and of cold
stratification on germination of H2SO4-scarified (for 1 h)
R. aromatica seeds following 4 weeks of incubation at
25/15oC in light. Data with the same superscript do not
differ statistically (Tukeys test, P 0.05).
Pretreatment
control
GA3, 2.5 mg/l
GA3, 12.5 mg/l
GA3, 25 mg/l
GA3, 500 mg/l
GA3, 1000 mg/l
7-day stratification
15-day stratification
25-day stratification

% germination after 4 weeks of

incubation at 25/15oC in light
16 1a
26 5b
43 9c
53 9c
92 3d
92 4d
89 2d
93 2d
98 1d

seeds collected at the same time but desiccated prior to

scarification and germination.
Physiological dormancy of R. aromatica seeds was
broken readily by either GA3 or stratification
treatments (Table 3). Compared to the control, a GA3

241

solution with a concentration as low as 2.5 mg/l

enhanced germination percentages significantly, from
16 1% to 26 5%, which increased further to 53 9%
at a GA3 concentration of 25 mg/l. At concentrations
of 500 and 1000 mg/l, GA3 completely alleviated
embryo dormancy, resulting in germination
percentages of 92 3% and 92 4%, respectively,
which were not statistically different than those of
seeds stratified for periods of 1 week to 25 days.
Once physical dormancy was broken, seeds of all
five species started to absorb water as soon as they were
placed on a moist substrate (Fig. 1). After 12 h of
imbibition, moisture content had increased to 52.0 5.9,
41.6 7.0, 43.2 6.7, 60.3 4.5, and 33.7 7.0% of the
initial seed weight in R. aromatica, R. glabra, R. trilobata,
R. typhina, and R. virens, respectively. By 24 h, seed
moisture content was 69.7 0.9% in R. aromatica,
61.4 1.4 % in R. glabra, 62.1 3.2% in R. trilobata,
72.4 0.9% in R. typhina, and 59.2 6.8% in R. virens.
After 48 h, seeds of all species visually had become fully
imbibed, with a moisture content of approx. 70-80%. By
72 h, the carpellary micropyle area (Li et al., 1999b) of
the endocarp in most seeds of R. typhina was beginning
to open, indicating that pressure was increasing in the

Figure 1. Water uptake (% of initial seed weight) during imbibition by scarified seeds of Rhus aromatica (j), R. glabra
(h), R. trilobata (n), R. typhina (s), and R. virens (d) (see Table 1 for details of scarification treatments). Essentially
no water (i.e., <3%) was taken up by non-scarified (control) seeds of any of the five species during the 1.0 h period.

242

X. Li et al.

radicle area; this did not occur in R. glabra, R. trilobata,

or R. virens until 96 h after the seeds were placed on
moist sand. Radicles of the four species began to
protrude 24 h after the appearance of this opening in
the carpellary micropyle area. This was coupled with
an obvious increase in moisture content. However,
no additional water was taken up by seeds of
R. aromatica after 48 h.
Neither temperatures nor light had any statistically
significant effects on germination of non-dormant
seeds of R. aromatica (Table 4). Germination reached
>90% in 4 weeks at all temperatures, including
15/6oC, in both light and darkness.
As in R. aromatica, light had no significant effect on
seed germination in R. glabra at any temperature. At
30/15oC, approx. 80% of the seeds germinated,
regardless of the light condition, i.e., white, red, farred, or constant darkness. However, germination in
this species is more sensitive to temperature than it is
in R. aromatica. Maximum germination (>90%) was
obtained at 25/15oC and at 20/10oC, which was
significantly higher than that (approx. 80%) obtained
at 30/15oC. Further, germination percentages at these
three temperature regimes were significantly higher
than those (approx. 70%) at either 35/20 or 15/6oC.
Nonetheless, germination percentages increased to
>90% at 15/6oC when the incubation period was
extended to 30 weeks, and it was not statistically
different from that at 25/15oC or at 20/10oC.
In R. trilobata, light also had no effect on seed
germination. Maximum germination was >90% in
both light and constant darkness at 30/15, 25/15, and
20/10oC after incubation for 4 weeks, and at 15/6oC
when seeds were incubated for 30 weeks. At
35/20oC, germination percentages decreased significantly to 80 2% in light and to 73 6% in darkness;
both of these percentages were considerably higher
than those of seeds at 15/6oC incubated in light
(58 8%) or in darkness (59 5%) for 4 weeks only.
Similarly, R. virens seeds germinated to high
percentages in light and in darkness at all five
temperature regimes. Maximum germination of 90%
was reached at 30/15, 25/15, and 20/10oC. At
35/20oC, germination percentages decreased to
79 8% in light and to 74 5% in darkness, which
were statistically comparable to those reached at
15/6oC. However, when incubation at 15/6oC was
extended to 30 weeks, germination percentages
increased to >90%, which was statistically equal to
germination of those at 25/15oC and at 20/10oC.

Discussion
Seeds of all seedlots of R. aromatica were
physiologically dormant and required pretreatments
for best germination results (Table 1), thus supporting

Table 4. Effects of light and of temperature on germination

of scarified (see Table 1 for details of treatments) seeds of
four Rhus species. For each species, data within a row
followed by the same uppercase superscript, or by the same
lowercase superscript within a column, do not differ
statistically (Tukeys test, P 0.05).
% germination after 4 weeks of
incubation in
Temperature (oC)

White light

Darkness

Rhus aromatica
35/20
30/15
25/15
20/10
15/6
15/62

95 3Aa
97 1Aa
98 1Aa
97 1Aa
94 2Aa
97 2Aa

89 3Aa
95 2Aa
97 1Aa
97 1Aa
92 2Aa
92 2Aa

Rhus glabra
35/20
30/153
25/15
20/10
15 /6
15/62

67 10Ac
82 1Ab
93 2Aa
95 2Aa
65 3Ac
92 1Aa

67 11Ac
81 3Ab
95 1Aa
92 1Aa
69 4Ac
90 1Aa

R. trilobata
35/20
30/15
25/15
20/10
15 /6
15/62

80 2Ab
94 2Aa
95 1Aa
95 3Aa
58 8Ac
97 1Aa

73 6Ab
90 1Aa
90 2Aa
91 2Aa
59 5Ac
94 1Aa

R. virens
35/20
30/15
25/15
20/10
15/6
15/62

79 8Ab
90 4Aab
95 4Aa
93 1Aa
75 5Ab
96 3Aa

74 5 Ab
89 4Aab
97 2Aa
93 1Aa
73 4Ab
92 3Aa

1
Seeds of this species were stratified at 5oC for 21 d prior to
incubation.
2
Final germination percentage at 15/6oC following 30 weeks
incubation after start of experiment.
3
Germination percentages at this temperature regime were
88 3 in red light and 88 1 in far-red light, which were not
different from that in white light or in darkness.

the conclusions of other investigators (Heit, 1967, 1970;

Brinkman, 1974; Young and Young, 1992).
Stratification at 5oC for only 7 days increased
germination to approx. 90%, which statistically was
comparable to that obtained by treating non-stratified
seeds with a GA3 solution of 500 or 1000 mg/l (Table
3). The fact that approx. 15% of the seeds of R.
aromatica (seedlot Fky94) germinated without
stratification and that GA3 effectively broke embryo
dormancy suggests that the physiological dormancy
involved here is non-deep (Baskin and Baskin, 1998).

Germination of Rhus seeds

Contrary to previous reports (Heit, 1967, 1970;

Brinkman, 1974; Young and Young, 1992) that seeds of
R. trilobata are physiologically dormant, seeds of the
two seedlots of this species used in the present study
germinated readily without pretreatment once
physical dormancy was broken (Table 1). Apparently,
this difference is due to seedlot variation (Heit, 1970).
However, it is interesting that the two seedlots, one
collected in Nebraska in 1996 and the other in Utah in
1994, behaved very similarly.
In accordance with reports in the literature (Lovell,
1964; Boyd, 1943; Farmer et al., 1982), scarified seeds of
all six seedlots of R. glabra germinated readily with or
without stratification (Table 1). Thus, dormancy in this
species is only caused by an impermeable fruit coat,
i.e., physical dormancy.
Almost all potentially viable seeds of R. typhina
germinated without stratification (Table 1), suggesting
that the embryo of this species is non-dormant.
Norton (1985, 1986, 1987) reported embryo dormancy
in R. typhina and tried various chemicals, including
GA3, to improve germination. However, the highest
germination obtained was 60%, compared to 20% in
control seeds. A possible reason for the low
germination percentages in the study may be that he
did not effectively scarify many of the seeds prior to
germination tests. Norton did not present data on
water uptake by the seeds.
No embryo dormancy was detected in R. virens
seeds. Up to 90% of the seeds germinated readily
without pretreatment (Table 1) in agreement with
Nokes (1986) and Hubbard (1986), who did not
recommend stratification to break seed dormancy in
this species. However, Tipton (1992) suggested a 75day stratification period for scarified seeds of R. virens
prior to incubation.
Maturation desiccation seems to be a prerequisite
for seeds of R. aromatica to respond to stratification
treatment (Table 2). That is, seeds require the terminal
drying stage of development to switch from a
developmental to a germinative mode. It is beyond the
scope of the present study to explain how this switch
works. However, we speculate that it might be related
to the ABA/GA3 balance in the seeds. It would be
interesting to test the effects of various concentrations
of GA3 on non-desiccated- and on desiccated seeds to
see if higher concentrations are required to stimulate
germination of non-desiccated embryos. However,
seeds of R. glabra and R. virens did not require
maturation desiccation prior to germination (Table 2),
which is required for germination of soybean
(Rosenberg and Rinne, 1986) and castor bean
(Kermode and Bewley, 1989). Embryos of mature
seeds of these four species are physiologically nondormant, whereas those of R. aromatica are
physiologically dormant (Table 1). This suggests that
maturation desiccation may be a requirement for

243

dormancy break in seeds with physiological

dormancy at maturity, but not for those that lack
physiological dormancy.
During imbibition, the seeds took up water
promptly and quickly. It only took 48 h for seeds of the
five species studied to become fully imbibed, i.e.,
moisture content of approx. 7080% of initial seed
weight (Fig. 1). This is very close to the moisture
content of fully imbibed seeds of R. ovata (approx.
75%) (Stone and Juhren, 1951). As a result of the
absence of embryo dormancy in seeds of R. glabra, R.
trilobata, R. typhina, and R. virens, full imbibition was
followed immediately by germination, which was
characterized by a continuous increase in moisture
content, resulting in the protrusion of the radicle. In
contrast, there was a long lag between full imbibition
and germination in seeds of R. aromatica, during which
no water was taken up by the seeds. Apparently, the
seeds were undergoing some internal physiological/
biochemical preparations for germination other than
cell growth/elongation, and thus they did not imbibe
water during this period. Our imbibition test was not
continued long enough for seeds of R. aromatica to
finish this physiological preparation process, though
given enough time, we predict that the moisture
content would increase after the termination of this
time-lag phase, resulting in protrusion of the radicle.
Neither presence nor absence of white light
significantly affected seed germination of any of the
four species studied (R. typhina not tested because of
insufficient seed supply) (Table 4). Further, R. glabra
seeds germinated equally well in white-, red- and farred light. Nonetheless, seeds usually germinated to a
slightly higher percentage in light than in darkness
(Table 4). Keeley (1987, 1991) reported that light
significantly increased germination of R. trilobata seeds
from California. Thus, we suggest that for best results,
light should be provided when conducting
germination tests for Rhus seeds, as recommended by
previous researchers (Lovell, 1964; Heit, 1967, 1970;
Brinkman, 1974; Li and Xu, 1989; Young and Young,
1992).
Overall, seed germination in the four sumac
species was rather insensitive to temperature, in
support of previous investigations on Rhus (Lovell,
1964; Heit, 1967; Brinkman, 1974; Farmer et al., 1982;
Washitani and Takenaka, 1986; Rasmussen and
Wright, 1988; Li and Xu, 1989; Tipton, 1992; Young and
Young, 1992). Highest germination percentages were
obtained at 30/15, 25/15, and 20/10oC in R. aromatica
and R. trilobata and at 25/15 and 20/10oC in R. glabra
and R. virens. As expected, the germination rate was
slow at 15/6oC, but given enough time the highest
final germination percentage occurred at this
temperature regime.
In summary, embryo dormancy was detected in
seeds of all four seedlots of R. aromatica, and it was

244

X. Li et al.

broken readily by either cold stratification or GA3.

Contrary to previous reports, seeds of R. trilobata
germinated well without any pretreatment, indicating
that the embryo is non-dormant. This also was the
case in seeds of R. glabra, R. typhina, and R. virens.
Maturation desiccation prior to cold stratification was
required for R. aromatica seeds to come out of
physiological dormancy, whereas non-desiccated
seeds of both R. glabra and R. virens germinated
readily. Fully imbibed seeds of the five species studied
have a moisture content of approx. 7080% of their
initial weight. Overall, neither light nor temperature
had significant effects on seed germination of the
species. These results agree with the generalization
that seeds with physical dormancy germinate under a
wide variety of environmental conditions once
physical dormancy is broken (Baskin and Baskin,
1998).

Acknowledgement
We thank T.E. Hemmerly, Middle Tennessee State
University, Murfreesboro, Drs. S. Meyer, USDA Shrub
Sciences Laboratory, Provo, Utah, and O.W. van
Auken, University of Texas at San Antonio, for
collecting fruits of Rhus aromatica, R. trilobata and
R. virens, respectively, used in the present study.

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Received 19 October 1998,

accepted after revision 31 March 1999
CAB International, 1999