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Seed Storage Behavior and Seed Germination of Nine Species of Lauraceae from Yunnan, China

云南九种樟科植物种子的萌发及脱水耐性



全 文 :云南九种樟科植物种子的萌发及脱水耐性
杨娅娟ꎬ 郭永杰ꎬ 秦少发ꎬ 胡枭剑∗
(中国科学院昆明植物研究所中国西南野生生物种质资源库ꎬ 昆明  650201)
摘要: 种子库是对野生植物进行长期迁地保存的重要手段ꎬ 但不适用于种子脱水敏感的植物ꎮ 樟科植物中
有不少为我国南方常见的重要经济林木ꎬ 许多种类具有较高的生态及经济价值ꎬ 但关于其种子萌发及脱水
耐性等生物学方面的研究资料十分缺乏ꎮ 本研究选取来源于 5个属的 9种樟科植物ꎬ 对其种子休眠及萌发
特性进行了初步的研究并利用 100粒种子法原理确定其脱水耐性ꎮ 结果表明樟树 (Cinnamomum camphora)
种子可能具有中度生理休眠ꎻ 毛尖树 (Actinodaphne forrestii)、 倒卵叶黄肉楠 (Actinodaphneo bovata)、 米槁
(Cinnamomum migao)、 网叶山胡椒 (Lindera metcalfiana var. dictyophylla)、 香叶树 (Lindera communis) 及多
果新木姜子 (Neolitsea polycarpa) 的种子具有浅生理休眠ꎻ 阴香 (Cinnamomum burmannii) 及粉叶楠 (Phoe ̄
be glaucophylla) 的种子可能不具有休眠ꎮ 全部 9种樟科植物种子在脱水到 2􀆰 86%~7􀆰 16%含水量后均失去全
部活力ꎻ 而保湿保存的种子含水量保持在 17􀆰 32%~44􀆰 87%之间ꎬ 仍保持全部或大部分活力ꎻ 因此该研究涉
及的 9种樟科植物都是脱水敏感性种子ꎬ 不适合传统的种子库保存方法ꎮ
关键词: 樟科ꎻ 种子ꎻ 休眠ꎻ 萌发ꎻ 脱水耐性
中图分类号: Q 945            文献标志码: A                文章编号: 2095-0845(2015)06-813-08
Seed Storage Behavior and Seed Germination of Nine
Species of Lauraceae from Yunnanꎬ China
YANG Ya ̄juanꎬ GUO Yong ̄jieꎬ QIN Shao ̄faꎬ HU Xiao ̄jian∗
(Germplasm Bank of Wild Species in Southwest of Chinaꎬ Kunming Institute of Botanyꎬ
Chinese Academy of Sciencesꎬ Kunming 650201ꎬ China)
Abstract: Seed banking following internationally agreed standards is an important way for preserving collections of
wild plant species ex situꎻ but this method is not suitable for desiccation sensitive species. Lauraceae comprehends
some of the dominant species in the evergreen broadleaved forest in the south of China and contains many species
both of ecological and economical importance. Howeverꎬ study on seed biology such as germination and desiccation
tolerance of this family is scarce. Seeds of 9 species from 5 genera of this family were collected and their dormancy
status and germination requirement were studiedꎻ also their desiccation tolerance were determined using a modified
100 ̄seed test. The results showed that seeds of Cinnamomum camphora probably have intermediate physiological dor ̄
mancyꎻ seeds of Actinodaphne forrestiiꎬ Actinodaphne obovataꎬ Cinnamomum migaoꎬ Lindera metcalfiana var. dictyo ̄
phyllaꎬ Lindera communis and Neolitsea polycarpa are non ̄deep physiological dormantꎻ Seeds of Cinnamomum bur ̄
mannii and Phoebe glaucophylla may have no or negligible dormancy. All 9 species lost seed viability after desiccated
to 2􀆰 86% - 7􀆰 16% moisture content while still retained considerable viability with moisture content ranged from
17􀆰 32% to 44􀆰 87% after moist storageꎻ thus seeds of the 9 species are all desiccation sensitive and can not be
stored at the conventional seed bank conditions.
Key words: Lauraceaeꎻ Seedꎻ Dormancy statusꎻ Germinationꎻ Desiccation tolerance
植 物 分 类 与 资 源 学 报  2015ꎬ 37 (6): 813~820
Plant Diversity and Resources                                    DOI: 10.7677 / ynzwyj201515099
∗ Author for correspondenceꎻ E ̄mail: huxiaojian@mail􀆰 kib􀆰 ac􀆰 cn
Received date: 2015-06-11ꎬ Accepted date: 2015-10-19
作者简介: 杨娅娟 (1986-) 女ꎬ 助理工程师ꎬ 主要从事种子脱水耐性ꎬ 储藏习性等相关研究及工作ꎮ
E ̄mail: yangyajuan@mail􀆰 kib􀆰 ac􀆰 cn
  Plant species in China as well as world wide
are facing greater threat nowadays than during the
recent geological past (Li and Pritchardꎬ 2009). To
counter this threatꎬ both in situ and ex situ conserva ̄
tion measurements should be taken into account. In
practiceꎬ seed banking following internationally a ̄
greed standards is mostly used to preserve collections
ex situ (Li and Pritchardꎬ 2009). Seed bank of the
Germplasm Bank of Wild Species ( GBOWS) in
Kunming China aiming on the conservation of wild
plant species of Chinaꎬ has preserved over 8 000
species with more than 30 000 collections by the end
of 2014. The conventional way for seed banking is
storage dry seeds (moisture content around 5%) at a
cold temperature ( around - 20 ℃). Howeverꎬ this
method is only effective for orthodox seeds which can
be dried without damageꎬ to low levels of moisture
contentꎬ and their longevity increases with decrease
in seed storage moisture content and temperature o ̄
ver a wide range (Robertsꎬ 1973)ꎻ for recalcitrant
and some intermediate seeds which are desiccation
sensitive to different extentꎬ other ways of long term
conservation such as cryopreservation have to be
considered. Thusꎬ determination of the seed desicca ̄
tion tolerance of target species is essential for suc ̄
cessful seed banking and ex situ conservation.
For determination of desiccation tolerance and
storage behaviorꎬ Hong and Ellis (1996) had deve ̄
loped a very precise methodꎬ but it costs thousands
of seeds. For tree species especially endangered tree
speciesꎬ it is difficult to collect enough seeds to con ̄
duct this experiment and often needs to compromise
the wild populationꎬ also the large ̄screening of des ̄
iccation tolerance requires considerable human re ̄
source and consumables ( Pritchard et al.ꎬ 2004).
Prichard et al. (2004) devised a less complicated
method to check the desiccation tolerance of several
palm species using only 100 seeds. The basic idea is
to compare the viability of both the freshly matured
seeds and the moist stored seeds with seeds after
desiccation. If the seeds after desiccation fail to ger ̄
minate while the fresh and moist stored seeds have
similar germination levelsꎬ the seeds are considered
to be desiccation sensitive. This method can not de ̄
termine species classed as intermediateꎬ but for the
seed bank (e􀆰 g. GBOWS) following the convention ̄
al seed storage protocolꎬ it is a very useful and eco ̄
nomical way for initial identification of desiccation
sensitive seeds.
Lauraceae comprehends some of the dominant
species in the evergreen broadleaved forest in the
southwest of China (Li and Pritchardꎬ 2009)ꎻ this
family is economically important as sources of medi ̄
cineꎬ timberꎬ nutritious fruits and perfumes ( Li et
al.ꎬ 2008). However its seed biology is under ̄re ̄
searched (Li and Pritchardꎬ 2009)ꎬ seed desicca ̄
tion tolerance of many species are still unknown.
Thus we collected 9 species from 5 genuses of this
family. For 8 of the 9 speciesꎬ seed germination re ̄
quirement and desiccation tolerance had not been re ̄
ported before.
To determine the seed desiccation toleranceꎬ
we adopted the basic principal of ‘100 ̄seeds test’
proposed by Prichard et al. (2004)ꎬ but as we had
some more seeds availableꎬ 3 replicates instead of 2
for the germination test were usedꎬ this allowed us to
perform an one ̄way ANOVA test for the germination
percentage data.
Before performing the ‘100 ̄seeds test’ꎬ it is de ̄
sirable to find an effective way for obtaining a relative ̄
ly high germination percentage as the indication of the
seed viability first. Thus we also conducted an experi ̄
ment to determine the temperature as well as hormone
requirement for seed germination of the 9 species.
1  Materials and methods
1􀆰 1  Seed collection
Nine species from 5 genera of Lauraceae were
collected during Novemberꎬ 2011ꎬ from different lo ̄
cations of Yunnan Provinceꎬ China (Table 1).
All collections were transported to the GBOWS
within 1 week. After cleaningꎬ all collections were
put in a room with a relative humidity (RH) of (75
± 10) % and a temperature of (15 ± 3) ℃ until the
418                                  植 物 分 类 与 资 源 学 报                            第 37卷
Table 1  Collection information of the 9 Lauraceae species
Species Date of collection           Locality Altitude / m
Actinodaphne forrestii 2011-11-6 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1450
A􀆰 bovata 2011-11-7 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1139
Cinnamomum burmannii 2011-11-7 Simaoꎬ Puerꎬ Yunnanꎬ China 1400
C􀆰 camphora 2011-11-7 Simaoꎬ Puerꎬ Yunnanꎬ China 1966
C􀆰 migao 2011-11-7 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1320
Lindera communis 2011-11-6 Xishanꎬ Kunmingꎬ Yunnanꎬ China 1450
L􀆰 metcalfiana var. dictyophylla 2011-11-3 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1400
Neolitsea polycarpa 2011-11-7 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1320
Phoebe glaucophylla 2011-11-6 Malipoꎬ Wenshanꎬ Yunnanꎬ China 1200
start of the experimentꎬ both of the desiccation treat ̄
ment and germination test started within 1 week.
Initial moisture content (MC) was determined
using 10 replicates with 1 seed for each replicate.
Fresh seeds were weighed by an electronic balance
(0􀆰 00001 g) and the dry weight was obtained using
the same balance after seeds were dried at 103 ℃ for
17 hours (ISTAꎬ 1999).
1􀆰 2  Germination of the freshly matured seeds
The germination test of the freshly matured seeds
was conducted under 20ꎬ 25ꎬ 30 and 25/ 10℃ (night /
day)ꎬ the photoperiod was 12 h light with 22􀆰 2 μmol􀅰
m-2s-1 illumination by cool white fluorescent light and
12 h dark. A gibberellic acid (GA3) treatment of 200
mg􀅰L-1 was also used for each temperature exclu ̄
ding 25 / 10 ℃ . In each treatmentꎬ 3 replicates of 25
seeds were used. The germination medium was 1%
agar / water or 1% agar / water with 200 mg􀅰L-1 GA3 .
The germination was checked every 7 days and a
seed with a radical more than 2 mm was considered
to be germinatedꎬ and the germination test was ter ̄
minated if no germination was recorded for consecu ̄
tive 4 weeks. At the point of terminationꎬ ungermi ̄
nated seeds were cut throughꎻ empty seeds were ex ̄
cluded for the calculation of germination percentage.
1􀆰 3  Desiccation and moist storage protocol
The desiccation of seeds were conducted by mix ̄
ing seeds with an equal weight of silica gel in a plas ̄
tic box sealed with an air ̄tight lid at 15 ℃. The equi ̄
librium relative humidity (eRH) of the seeds were
measured every 2 days using a Rotronic HC2 ̄AW
probe attached to a HygroLab C1 unit (Rotronic Ltd.ꎬ
Crawleyꎬ UK)ꎬ the silica gel was changed after the
measurement of eRHꎬ once it equaled to or below
15%ꎬ the desiccation process was terminated and
the moisture content was determined using the same
method as the determination of initial moisture con ̄
tent. Also 3 replicates of 25 seeds were used for the
germination test with the best temperature and hor ̄
mone combination obtained from the germination test
of freshly matured seeds mentioned above.
Once the desiccation process finishedꎬ seeds of
the same species moist stored were also taken out for
MC determination and germination test which follow
the same protocol of the desiccated seeds.
1􀆰 4  Statistical analysis
Moisture content (MC) was calculated as:
MC=(FW ̄DW) / FW
Where FW is the fresh weight and DW is the
dry weight.
Germination percentage (GP) and mean germi ̄
nation time (MGT) was calculated as:
GP =∑ni / N
MGT(days)= ∑(ti∗ni) / ∑ni
Where ti is the number of days from experiment
startingꎬ ni is the number of seeds germinated at
each checking day and N is the total number of
seeds tested.
For freshly matured seedsꎬ we analyzed the ger ̄
mination percentages and mean germination time u ̄
sing a univariate General Linear Model to test the
effect of temperatureꎻ a Student ̄Newman ̄Keuls (S ̄
N ̄K) post ̄hoc test was applied for multiple compari ̄
sons between different temperaturesꎻ while a T test
5186期      YANG Ya ̄juan et al.: Seed Storage Behavior and Seed Germination of Nine Species of Lauraceae from 􀆺     
was applied for the effect of GA3 at each temperature
excluding 25 / 10 ℃ .
For seeds that desiccated and moist storedꎬ we
compared the germination percentage with the initial
germination percentage of the freshly matured seeds
by applying an one way ANOVA with a Student ̄New ̄
man ̄Keuls (S ̄N ̄K) post ̄hoc test.
The percentage data were arcsine transformed
before analysis. Differences obtained at a level of P<
0􀆰 05 were considered to be significant. All statistical
analysis was carried out by SPSS 16􀆰 0 ( Chicagoꎬ
ILꎬ USA).
2  Results
2􀆰 1  Moisture content
Ranges of the moisture contents of the freshly
matured seedsꎬ seeds after desiccation and seeds af ̄
ter moist storage are 14% to 45%ꎬ 2􀆰 86% to 7􀆰 16%
and 17􀆰 32% to 44􀆰 87% respectively (Table 2).
2􀆰 2  Germination of freshly matured seeds
Five of total 9 species achieved a relatively high
germination percentage ( over 80%) of freshly ma ̄
tured seedsꎻ while 2 species germinated to 60% -
70% and the other 2 species germinated around
45%. Though most of them germinated readily at 20
or 25 / 10 ℃ꎬ the optimal germination temperature
varied among different species (Table 3).
The effect of GA3 also varied among different
speciesꎬ for it promoted GP and reduced MGT both
significantly for seeds of C􀆰 migao and L􀆰 metcalfiana
var. dictyophyllaꎬ while no significant effect was shown
on GP or MGT on C􀆰 camphora and P􀆰 glaucophylla.
For A􀆰 obovata and L􀆰 communis GA3 only promoted
germination percentage at some temperatures while for
N􀆰 polycarpa only MGT was reduced significantly by
GA3 at 20 ℃. There was also a significantly negative
effect of GA3 on C􀆰 burmannii at 25 ℃ꎬ and MGT was
slightly reduced by GA3 at 20 ℃ (Table 3 and 4).
2􀆰 3  Response to desiccation and moist storage
Moist stored seeds of the 2 Actinodaphne species
and L􀆰 metcalfiana var. dictyophylla germinated equal ̄
ly to freshly matured seeds while no seed germinated
after desiccation. The rest 6 species showed signifi ̄
cant reduction after moist storageꎬ but still retained
the ability to germinate while no germination occurred
after desiccation (Table 2).
3  Discussion
Though there was a significant reduction of MGT
by GA3 at 20 ℃ for seeds of C􀆰 burmanniiꎬ but the
germination was checked every 7 daysꎬ the difference
between the two MGT data was within 7 daysꎬ so
this difference could be due to systematic error and
had no significant biological meaningꎻ thus we specu ̄
lated that seeds of C􀆰 burmannii and P􀆰 glaucophylla
may have no or negligible dormancyꎬ for they germi ̄
nated readily at some temperatures and GA3 had no
effect on either GP or MGT. Seeds of C􀆰 camphora
Table 2  Effects of desiccation and moist storage on seed germination of the 9 Lauraceae species
Seed lot
Fresh
Moisture
content / %
Germination
/ %
Desiccation
Moisture
content / %
Germination
/ %
Moist storage
Moisture
content / %
Germination
/ %
Actinodaphne forrestii 43􀆰 54 ± 5􀆰 89 93􀆰 33 ± 11􀆰 55a 5􀆰 73 ± 1􀆰 20 0c 35􀆰 00 ± 2􀆰 09 86􀆰 67 ± 8􀆰 32a
A􀆰 obovata 45􀆰 85 ± 4􀆰 73 84􀆰 33 ± 15􀆰 01a 3􀆰 49 ± 0􀆰 35 0c 44􀆰 87 ± 7􀆰 01 77􀆰 67 ± 4􀆰 04a
Cinnamomum burmannii 25􀆰 85 ± 1􀆰 14 96􀆰 00 ± 4􀆰 00a 3􀆰 69 ± 1􀆰 03 0c 17􀆰 32 ± 1􀆰 68 64􀆰 00 ± 4􀆰 00b
C􀆰 camphora 26􀆰 32 ± 1􀆰 08 70􀆰 67 ± 4􀆰 62a 3􀆰 56 ± 1􀆰 08 0c 21􀆰 30 ± 1􀆰 35 45􀆰 33 ± 2􀆰 31b
C􀆰 migao 35􀆰 02 ± 4􀆰 41 60􀆰 00 ± 8􀆰 00a 2􀆰 86 ± 0􀆰 27 0c 19􀆰 52 ± 7􀆰 04 49􀆰 33 ± 8􀆰 33b
Lindera communis 23􀆰 24 ± 11􀆰 45 44􀆰 00 ± 4􀆰 00a 3􀆰 69 ± 0􀆰 85 0c 23􀆰 61 ± 8􀆰 86 29􀆰 33 ± 2􀆰 31b
L􀆰 metcalfiana var. dictyophylla 24􀆰 79 ± 5􀆰 73 48􀆰 00 ± 22􀆰 27a 6􀆰 31 ± 3􀆰 81 0c 26􀆰 56 ± 11􀆰 87 30􀆰 66 ± 12􀆰 22a
Neolitsea polycarpa 24􀆰 04 ± 6􀆰 86 80􀆰 00 ± 10􀆰 58a 2􀆰 87 ± 0􀆰 32 0c 19􀆰 52 ± 7􀆰 04 52􀆰 00 ± 10􀆰 58b
Phoebe glaucophylla 37􀆰 99 ± 3􀆰 56 92􀆰 00 ± 10􀆰 58a 7􀆰 16 ± 0􀆰 45 0c 43􀆰 57 ± 7􀆰 53 68􀆰 00 ± 6􀆰 93b
Data are means ± standard deviation. Different letters represent significant different treatment means (within the same species) by S ̄N ̄K test at 5%
level of significance
618                                  植 物 分 类 与 资 源 学 报                            第 37卷
Table 3  Effects of different treatments on germination percentage (%) of freshly matured seeds
of 9 Lauraceae species at different temperatures
Species Temprature / ℃
GA3 solution concentration / mg􀅰L-1
0 200
Total by temprature
Actinodaphne forrestii 20   33􀆰 33 ± 6􀆰 11   49􀆰 33 ± 10􀆰 06   41􀆰 33 ± 11􀆰 50a
25   77􀆰 33 ± 8􀆰 33   90􀆰 67 ± 10􀆰 07   84􀆰 00 ± 11􀆰 03b
30   28􀆰 00 ± 2􀆰 00   58􀆰 76 ± 10􀆰 06   43􀆰 33 ± 21􀆰 97a
25 / 10   93􀆰 33 ± 11􀆰 55       —   93􀆰 33 ± 11􀆰 55b
Total by treatment   58􀆰 00 ± 31􀆰 15   66􀆰 22 ± 20􀆰 70       —
A􀆰 obovata 20   51􀆰 00 ± 10􀆰 15   73􀆰 33 ± 6􀆰 51∗   62􀆰 17 ± 14􀆰 41ab
25   66􀆰 67 ± 23􀆰 71   84􀆰 33 ± 15􀆰 01   75􀆰 50 ± 20􀆰 22b
30   17􀆰 67 ± 30􀆰 60   46􀆰 67 ± 43􀆰 84   32􀆰 17 ± 37􀆰 36ac
25 / 10       0       —       0c
Total by treatment   33􀆰 83 ± 32􀆰 39   68􀆰 11 ± 28􀆰 79       —
Cinnamomum burmannii 20   96􀆰 00 ± 4􀆰 00   96􀆰 00 ± 4􀆰 00   96􀆰 00 ± 3􀆰 58a
25   90􀆰 67 ± 2􀆰 31   70􀆰 67 ± 4􀆰 62∗   80􀆰 67 ± 11􀆰 43b
30       0       0       0c
25 / 10       0       —       0c
Total by treatment   46􀆰 67 ± 48􀆰 82   55􀆰 56 ± 43􀆰 19       —
C􀆰 camphora 20       0   2􀆰 67 ± 2􀆰 31   1􀆰 33 ± 2􀆰 07a
25   1􀆰 33 ± 2􀆰 31   2􀆰 67 ± 2􀆰 31   2􀆰 00 ± 2􀆰 19a
30   17􀆰 33 ± 2􀆰 31   21􀆰 33 ± 12􀆰 86   19􀆰 33 ± 8􀆰 55b
25 / 10   70􀆰 67 ± 4􀆰 62       —   70􀆰 67 ± 4􀆰 62c
Total by treatment   16􀆰 57 ± 24􀆰 44   8􀆰 89 ± 11􀆰 45       —
C􀆰 migao 20   24􀆰 00 ± 16􀆰 00   29􀆰 33 ± 2􀆰 31   26􀆰 67 ± 10􀆰 63a
25   12􀆰 00 ± 6􀆰 93   32􀆰 00 ± 10􀆰 58   22􀆰 00 ± 13􀆰 56a
30   5􀆰 33 ± 2􀆰 31   60􀆰 00 ± 8􀆰 00∗   32􀆰 67 ± 30􀆰 40a
25 / 10   28􀆰 00 ± 8􀆰 00       —   28􀆰 00 ± 8􀆰 00a
Total by treatment   17􀆰 33 ± 12􀆰 57   40􀆰 44 ± 16􀆰 18       —
Lindera communis 20   0􀆰 08 ± 0􀆰 00   44􀆰 00 ± 4􀆰 00∗   26􀆰 00 ± 19􀆰 88a
25   12􀆰 00 ± 6􀆰 93   22􀆰 67 ± 15􀆰 14   17􀆰 33 ± 12􀆰 04a
30       0       0       0b
25 / 10       0       —       0b
Total by treatment   5􀆰 00 ± 6􀆰 18   22􀆰 22 ± 20􀆰 60       —
L􀆰 metcalfiana var. dictyophylla 20   33􀆰 33 ± 16􀆰 65   48􀆰 00 ± 22􀆰 27   40􀆰 67 ± 19􀆰 34a
25   5􀆰 33 ± 4􀆰 62   14􀆰 67 ± 4􀆰 62   10􀆰 00 ± 6􀆰 57b
30       0   10􀆰 67 ± 2􀆰 31∗   5􀆰 33 ± 6􀆰 02b
25 / 10   12􀆰 00 ± 6􀆰 93       —   12􀆰 00 ± 6􀆰 93b
Total by treatment   12􀆰 67 ± 15􀆰 43   24􀆰 44 ± 21􀆰 11       —
Neolitsea polycarpa 20   80􀆰 00 ± 10􀆰 58   77􀆰 33 ± 14􀆰 05   78􀆰 67 ± 11􀆰 22a
25   40􀆰 00 ± 14􀆰 42   36􀆰 00 ± 0􀆰 00   38􀆰 00 ± 9􀆰 38b
30       0   4􀆰 00 ± 6􀆰 93   2􀆰 00 ± 4􀆰 90c
25 / 10   45􀆰 33 ± 22􀆰 03       —   45􀆰 33 ± 22􀆰 03b
Total by treatment   41􀆰 33 ± 32􀆰 01   39􀆰 11 ± 32􀆰 79       —
Phoebe glaucophylla 20   72􀆰 00 ± 6􀆰 93   74􀆰 67 ± 12􀆰 22   73􀆰 33 ± 9􀆰 04a
25   54􀆰 67 ± 4􀆰 62   50􀆰 7 ± 6􀆰 11   52􀆰 67 ± 5􀆰 31b
30   12􀆰 00 ± 12􀆰 00   26􀆰 67 ± 12􀆰 86   19􀆰 33 ± 13􀆰 72c
25 / 10   92􀆰 00 ± 10􀆰 58       —   92􀆰 00 ± 10􀆰 58d
Total by treatment   57􀆰 67 ± 31􀆰 75   50􀆰 67 ± 22􀆰 80       —
Data are means ± standard deviation. Asterisk (∗) represents the significant effect of GA3 treatment. Different letters represent significant different
temperature means (within the same species) by S ̄N ̄K test at 5% level of significance. Dash (—) represents data unavailable
7186期      YANG Ya ̄juan et al.: Seed Storage Behavior and Seed Germination of Nine Species of Lauraceae from 􀆺     
Table 4  Effects of different treatments on mean germination time (days) of freshly matured seeds
of the 9 Lauraceae species at different temperatures
Species Temprature / ℃
GA3 solution concentration / mg􀅰L-1
0 200
Total by temprature
Actinodaphne forrestii 20     155􀆰 33 ± 22􀆰 68     65􀆰 14 ± 8􀆰 66∗   110􀆰 23 ± 51􀆰 73a
25     95􀆰 01 ± 10􀆰 89     41􀆰 44 ± 0􀆰 74∗   68􀆰 22 ± 30􀆰 14b
30     58􀆰 50 ± 9􀆰 96     44􀆰 38 ± 3􀆰 76   51􀆰 44 ± 10􀆰 25c
25 / 10     85􀆰 49 ± 0􀆰 70     —   85􀆰 49 ± 0􀆰 70d
Total by treatment     98􀆰 58 ± 38􀆰 73     50􀆰 32 ± 12􀆰 15   —
A􀆰 obovata 20     108􀆰 76 ± 16􀆰 74     104􀆰 58 ± 27􀆰 90   106􀆰 67 ± 20􀆰 70a
25     100􀆰 36 ± 60􀆰 18     54􀆰 17 ± 5􀆰 25   77􀆰 27 ± 45􀆰 83ab
30     28􀆰 00 ± 48􀆰 50     33􀆰 79 ± 29􀆰 26   30􀆰 89 ± 35􀆰 96bc
25 / 10     —     —   —
Total by treatment     59􀆰 28 ± 59􀆰 08     64􀆰 18 ± 37􀆰 57   —
Cinnamomum burmannii 20     38􀆰 12 ± 0􀆰 81     32􀆰 19 ± 0􀆰 68∗   35􀆰 15 ± 3􀆰 32a
25     27􀆰 68 ± 1􀆰 10     29􀆰 26 ± 2􀆰 54   28􀆰 47 ± 1􀆰 95b
30     —     —   —
25 / 10     —     —   —
Total by treatment     16􀆰 45 ± 17􀆰 62     20􀆰 48 ± 15􀆰 47   —
C􀆰 camphora 20     —     14􀆰 00 ± 12􀆰 12   7􀆰 00 ± 10􀆰 84a
25     9􀆰 33 ± 16􀆰 16     18􀆰 67 ± 16􀆰 16   14􀆰 00 ± 15􀆰 34a
30     35􀆰 93 ± 1􀆰 61     43􀆰 68 ± 7􀆰 64   39􀆰 81 ± 6􀆰 51b
25 / 10     136􀆰 97 ± 14􀆰 64     —   136􀆰 97 ± 14􀆰 64c
Total by treatment     45􀆰 56 ± 57􀆰 58     25􀆰 45 ± 17􀆰 54   —
C􀆰 migao 20     122􀆰 42 ± 41􀆰 00     50􀆰 12 ± 12􀆰 60∗   57􀆰 71 ± 57􀆰 23a
25     33􀆰 73 ± 5􀆰 47     28􀆰 19 ± 1􀆰 47   30􀆰 96 ± 4􀆰 69b
30     71􀆰 17 ± 10􀆰 10     84􀆰 25 ± 30􀆰 64   77􀆰 71 ± 21􀆰 63a
25 / 10     137􀆰 33 ± 10􀆰 60     —   137􀆰 33 ± 10􀆰 60c
Total by treatment     89􀆰 78 ± 48􀆰 80     56􀆰 03 ± 27􀆰 93   —
Lindera communis 20     63􀆰 00 ± 14􀆰 00     102􀆰 83 ± 8􀆰 23∗   82􀆰 92 ± 24􀆰 11a
25     36􀆰 40 ± 13􀆰 50     53􀆰 28 ± 5􀆰 51   44􀆰 84 ± 13􀆰 06b
30     —     —   —
25 / 10     —     —   —
Total by treatment     24􀆰 85 ± 28􀆰 96     52􀆰 04 ± 44􀆰 81   —
L􀆰 metcalfiana var. dictyophylla 20     85􀆰 63 ± 9􀆰 86     54􀆰 08 ± 13􀆰 82∗   69􀆰 86 ± 20􀆰 34a
25     33􀆰 83 ± 31􀆰 76     35􀆰 00 ± 7􀆰 00   34􀆰 42 ± 20􀆰 58b
30     —     28􀆰 00 ± 0􀆰 00∗   14􀆰 00 ± 15􀆰 34c
25 / 10     44􀆰 10 ± 5􀆰 28     —   44􀆰 10 ± 5􀆰 28b
Total by treatment     40􀆰 89 ± 34􀆰 99     39􀆰 03 ± 14􀆰 02   —
Neolitsea polycarpa 20     150􀆰 57 ± 3􀆰 24     118􀆰 58 ± 4􀆰 19∗   134􀆰 58 ± 17􀆰 84a
25     88􀆰 35 ± 7􀆰 23     75􀆰 70 ± 8􀆰 57   82􀆰 03 ± 9􀆰 91b
30     —     18􀆰 67 ± 32􀆰 33   9􀆰 33 ± 22􀆰 86c
25 / 10     135􀆰 15 ± 13􀆰 97     —   135􀆰 15 ± 13􀆰 97a
Total by treatment     93􀆰 52 ± 61􀆰 64     70􀆰 98 ± 46􀆰 57   —
Phoebe glaucophylla 20     57􀆰 15 ± 5􀆰 86     52􀆰 41 ± 3􀆰 09   54􀆰 78 ± 4􀆰 93a
25     42􀆰 23 ± 3􀆰 13     39􀆰 02 ± 7􀆰 75   40􀆰 62 ± 5􀆰 57b
30     19􀆰 05 ± 16􀆰 51     30􀆰 11 ± 5􀆰 05   24􀆰 58 ± 12􀆰 49c
25 / 10     81􀆰 93 ± 3􀆰 64     —   81􀆰 93 ± 3􀆰 64d
Total by treatment     50􀆰 09 ± 25􀆰 09     40􀆰 51 ± 10􀆰 88   —
Data are means ± standard deviation. Asterisk (∗) represents the significant effect of GA3 treatment. Different letters represent significant different
temperature means (within the same species) by S ̄N ̄K test at 5% level of significance. Dash (—) represents data unavailable
818                                  植 物 分 类 与 资 源 学 报                            第 37卷
was reported to be physiological dormantꎬ and need
cold stratification for 4 months as well as a H2 O2
treatment ( Chien and Linꎬ 1999)ꎻ in this studyꎬ
seeds have no response to GA3ꎬ so we speculated
that they were intermediate dormant according to the
classification system of Baskin and Baskin (2004).
For the rest 6 speciesꎬ due to their positive response
to GA3ꎬ we infer that they have non ̄deep physiologi ̄
cal dormancy (Baskin and Baskinꎬ 2004). Howev ̄
erꎬ determination of dormancy type was not the main
purpose of this studyꎬ the primary aim for conduc ̄
ting germination experiment was to obtain an accept ̄
able germination as a parameter for the testing of
seed viabilityꎻ more detailed work still need to be
done for the 2 Lindera species and C􀆰 migao to opti ̄
mize the germination.
To dateꎬ only one Cassytha species from Aus ̄
tralia is confirmed to be desiccation tolerant in the
family of Lauraceae (Royal Botanic Gardens Kewꎬ
2015)ꎻ and different genera are unevenly studied.
For genera of Actinodaphne and Phoebeꎬ no data is
available for any species. Our data on the 2 species
from Actinodaphne and 1 species of Phoebe showed
all of them are desiccation sensitive. Cinnamomum is
the most intensively studied genus of this family and
3 species were confirmed to be recalcitrant (Royal
Botanic Gardens Kewꎬ 2015). Probably due to its
relatively wide distribution and economical impor ̄
tanceꎬ C􀆰 camphora is one of the most studied spe ̄
cies of the family. Different researches implied dif ̄
ferent desiccation tolerance. Study on collection from
Nepal showed seeds still retained some germinibility
after dry stored at ambient temperatures for 6 or 12
monthsꎬ but exact moisture content was not stated
(Campbellꎬ 1980). Chien and Lin (1999) desicca ̄
ted seeds collected from Taiwan province to 6􀆰 7%ꎬ
a large portion of the desiccated seeds survived after
12 months of storage at 5 ℃ or 15 ℃ but lost viabili ̄
ty after 1 month at -20 ℃ꎬ thus they classified the
seeds of C􀆰 camphora as intermediate. In this studyꎬ
moisture content was reduced to 3􀆰 56% and all
seeds lost viability. The different response could be
due to different levels of desiccationꎬ howeverꎬ ori ̄
gin of collection could also be a factor (Hong and
Ellisꎬ 1996). Similar result was shown for Camellia
sinensisꎬ desiccation tolerance differed between col ̄
lections from different sitesꎬ but non of them survived
-20 ℃ (Chen et al.ꎬ 2012). Though the exact stor ̄
age behavior of this specie is still unclearꎬ based on
the current dataꎬ we consider seeds of C􀆰 camphora
can not be stored under the conventional seed bank
condition (i􀆰 e. 5% MC at -20 ℃).
For the rest of the two Cinamomum species
studied hereꎬ significant reduction were shown on
GP after moist storageꎬ we speculated the loss of via ̄
bility could be due to the reduction of the moisture
contentꎬ and loss of viability with time during moist
storage could also be a reasonꎻ howeverꎬ more than
45% of seeds still retained their viability whilst no
germination occurred after desiccated to a MC below
5%ꎻ these two species can be classified as desicca ̄
tion sensitive.
The two species from genus Lindera in this study
are also desiccation sensitiveꎬ another species L􀆰 meg ̄
aphylla is classified as intermediate (Royal Botanic
Gardens Kewꎬ 2015). For the two species of Neolit ̄
sea with known storage behaviorꎬ one is classified as
recalcitrant and another intermediate (Royal Botanic
Gardens Kewꎬ 2015). In this studyꎬ seeds of N􀆰 poly ̄
carpa showed a significant reduction after moist stor ̄
ageꎬ but still retained more than 50% germinibility
while no germination occurred after desiccation to
2􀆰 87% MCꎻ thus we consider this species to be des ̄
iccation sensitive.
Though the result of this research did not fit
perfectly into the 3 categories showed in the article
of Prichard et al. ( 2004)ꎬ the effect of moisture
content on seed viability can be distinguishedꎬ for
all the 9 species of Lauraceaeꎬ no germination oc ̄
curred after desiccation while still considerable ger ̄
minability retained after moist storage of the same
period. According to the principle of ‘100 ̄seeds test’ꎬ
all these 9 species of Lauraceae can be classified as
desiccation sensitive.
9186期      YANG Ya ̄juan et al.: Seed Storage Behavior and Seed Germination of Nine Species of Lauraceae from 􀆺     
Based on the current data on seed desiccation
tolerance of Lauraceaeꎬ only one species from the
genus Cassytha L. was confirmed to be desiccation
tolerantꎬ and species of this genus are different in
being parasitic vines with other tree of shrub species
of this family. Only one Cassytha species distributed
in Chinaꎬ and its desiccation tolerance is unknown.
As this family is both ecologically and economically
importantꎬ more detailed work should be done on its
seed biology in the future.
In conclusionꎬ seeds of C􀆰 burmannii and P􀆰 glau ̄
cophylla may have no or negligible dormancyꎬ seeds
of C􀆰 camphora may be intermediate physiological
dormantꎬ while seeds of A􀆰 forrestiiꎬ A􀆰 obovataꎬ C􀆰 mi ̄
goꎬ L􀆰 communisꎬ L􀆰 metcalfiana var. dictyophylla and
P􀆰 glaucophylla are probably non ̄deep physiological
dormant. Seeds of the 9 Lauraceae species are all
discussion sensitiveꎬ thus can not be preserved by
the conventional seed bankꎬ other methods for long
term conservation ex situ such as cryo ̄preservation
should be considered.
Acknowledgements: We convey our great thanks to Profes ̄
sor Hugh Prichardꎬ Professor YANG Xiang ̄yun and Doctor LI
Ai ̄hua for their advice and help on experimental design. We
also thank GUO Yun ̄gang for his help with the experiment.
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028                                  植 物 分 类 与 资 源 学 报                            第 37卷