Download Seedling establishment in Scaligeria stewartiana (Nasir) Nasir

ISSN 2449-8955
European Journal
of Biological Research
Research Article
Seedling establishment in Scaligeria stewartiana (Nasir)
Nasir (Apiaceae) from North-West Himalayas, India
Kulvinder Kour1, B. L. Bhellum1*, I. A. Hamal2
1
Department of Botany, Govt. College for Woman Parade, Jammu - 180001, Jammu & Kashmir State, India
BGSB University, Rajouri, Jammu & Kashmir State, India
*Corresponding author: B. L. Bhellum; E-mail: [email protected]
2
Received: 25 April 2016; Revised submission: 10 June 2016; Accepted: 21 June 2016
Copyright: © The Author(s) 2016. European Journal of Biological Research © T.M.Karpiński 2016. This is an open access article
licensed under the terms of the Creative Commons Attribution Non-Commercial 4.0 International License, which permits
unrestricted, non-commercial use, distribution and reproduction in any medium, provided the work is properly cited.
DOI: http://dx.doi.org/10.5281/zenodo.56106
ABSTRACT
1. INTRODUCTION
The present paper deals with the morphological
diversity of seedling establishment in different
populations of Scaligeria stewartiana (Nasir) Nasir,
inhabiting in North-West Himalayas. The seedling
at different stages in the population of the species
revealed marked variability. Seed treatments for
germination resulted in understanding of seed
dormancy and time required for the seedling
establishment. The species develops variability in
the genetic level that is transmitted and reflected in
the form of morphology in order to adapt itself in
the extremes of biotic and abiotic variables.
Photographs of the seedling and data compiled in
tables catch the specific stages of their seedling
development after germination as well variability of
the chromosomes. The initiation of tuber formation
under the ground level in the soil and allocation of
resources perform indispensable role in the life
span as well as in paving the way for survival,
conservation and distribution of the species.
Scaligeria stewartiana (Nasir) Nasir (Tribe
pyramidoptereae, Apiaceae) has undergone numerous nomenclatural changes both at genus and
species level. Currently, the number of species has
been reduced to three. The adaptation of living
organisms to the ecological amplitude of any habitat
depends upon their genetic composition, which in
turn is controlled by various components of their
genetic system and finally translated and expressed
in the form of morphology. Members of the family
Apiaceae have exploited several strategies in order
to achieve tremendous diversity in their habitat and
geographical distribution in the Himalayas that
support rich umbellifer diversity [1]. Scaligeria
stewartiana (Nasir) Nasir was recorded from
Shiwaliks in Jammu region as a new record for
India [2]. The species inhabits isolated populations
and turned out to be a diploid, having 2n=20 besides
carrying 1-4 extra chromosomes [3]. It is locally
known as Jangli Ajwain, the fruit of the species is
used as carminative and spice. Plant has thrived
high for its conservation by naturally allocating
most of its resources towards below ground part,
tuber for its survival. Another threat towards its
diminishing population is its long period of seedling
establishment which takes about four years to establish and a seed dormancy period of full one year.
Keywords: Seed germination; Seedlings; Seed set;
Tuber; Scaligeria stewartiana; India.
European Journal of Biological Research 2016; 6 (3): 145-151
146 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
The period of production of seeds year after
year is also restricted exactly to four years in which
the environmental disturbances, effect of local activities has imposed a lot on its survival. The seeds as
well as the leaves are used for various types of
gastrointestinal disorders by the locals. The seeds
are also used for treatment of digestive disorders of
live-stock.
2. MATERIALS AND METHODS
Scaligeria stewartiana (Nasir) Nasir of the
family Apiaceae is growing in wild situations of
different parts of Jammu Province expanding to the
different locations at a distance of over 100 Kms,
ranging in altitude from 600 to 1200 mts. The
sources of the tubers are confined to different
localities namely Mansar, Tikri and Reasi. The
tubers were transplanted in the botanical garden.
Seeds were collected from the different populations
and were treated for their germination. Seed
germination has been studied under laboratory as
well as in the experimental plots of the University
Botanic Garden. The seeds collected during May
1995, were treated with 0.1% mercuric chloride for
two minutes. Thereafter, the seeds were given a
wash in 70 percent alcohol. All traces of mercuric
chloride and alcohol were removed by washing
the seeds thoroughly with distilled water. For
germination in the laboratory, the seeds were
divided into replicates of 50 seeds each and then
subjected to following physical and chemical
treatments.
a) Chilling at ± 50C for 15, 20, 30 and 50 days using
refrigerator
b) Treatment with conc. sulphuric acid for two
minutes.
c) Hard water treatment for 2 minute.
d) Seed coat puncturing
e) Seed coat scarification.
In all the above treatments including control,
no seed germination was recorded under laboratory
conditions. However, from the seeds collected
during May, 1995, 53 seeds germinated in the
experimental plots of the Botanic Garden during
January 1997, after sowing during October 1995.
There was no germination of the seeds from October
1995 to December 1996. The germinated seeds
formed well-established seedlings and some of these
seedlings developed into mature plants and flowered
in the fourth year. The critical period starts even
before germination with predation of seeds,
followed by the germination phase that usually
allows only less than half of the viable seed to
viable seed to emerge, predation on germination and
seedlings.[4]
Seed germinator was also used in order to
germinate seeds. However seeds did not respond to
the different treatments of variable temperature
provided to facilitate germination. Observation of
Botanical Garden population where tubers were
transplanted three years earlier, seedlings were
observed growing. The measurements of seedlings
were made and data was collected. All photos were
done by using Olympus PM6 35 Camera.
3. RESULTS
3.1. Observations
The seeds from natural population growing at
Mansar were collected and treated for germination
and showed dormancy of one year (Fig. 1). The
germination of seed after dormancy resulted in the
formation of radical, which is first to emerge and
grow geotropically. After the emergence of the
radical two cotyledons are formed which on
reaching above ground start turning green. Both the
cotyledons were unequal in size in majority of the
seedlings and remain green for 20 to 30 days.
Seedlings attained a length of 5.0-5.4 cm and the
primary root also grow in size which varies from
3.0-3.4 cm. The elongation of primary root resulted
in initiation of a tuber in the form of a small
swelling at the sub apical position. However, there
is no shoot formation. At the end of the growing
season the cotyledons wither and the seedlings
perennate through the tuber. The epidermis of the
tuber starts turning light brown. As the tuber sprouts
during next growing period, a basal leaf emerges
from tuber above ground (Fig. 1 and Fig. 2 A-B).
The basal leaf of the second year seedlings
remained green for 30-40 days. During this period
photosynthesis in leaves provide nutrition to
developing tubers, which attained a size of 8-9.5
mm. The leaves again withered towards the end of
the growing season, however, tuber perennated in
the soil and it penetrates in the soil up to 15 cm deep
European Journal of Biological Research 2016; 6 (3): 145-151
147 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
below the ground level. The tubers attain almost the
normal size and perennate for the next year (Fig. 2
C). During third year growing season, the tuber
sprouted again which resulted in the formation of
2-3 basal leaves as shown in (Fig. 2 D). After
further growth the basal leaves emerged above the
soil and turned green. The mature leaves formed
during third year are larger in size more dissected
and remained above ground for 35-45 days. All the
resources gathered during growing season were
allocated towards below ground tuber [5].
Figure 3. Scaligeria stewartiana (Nasir) Nasir: a matured
tuber exposed to show under the ground level.
Figure 1. Population of Scaligeria stewartiana (Nasir)
Nasir in Mansar.
Figure 2. Photographs of seedling of Scaligeria
stewartiana (Nasir) Nasir showing different stages A-C:
Seedlings at first year after seed germination; D - Second
year seedling showing one basal leaf arising from the
tuber; E - Third year seedling showing more than one
leaf; F - Fourth year tuber showing numerous basal
leaves above ground.
The tuber that was rounded in the first and
second year of growing season attains irregular
shape. The leaves after the completion of the
growing season wither and tubers enter into
dormancy. The tuber which is irregular in shape
attaining brownish to black colour in its outer
covering again perennates for fourth year and
sprouted in the forthcoming season of growth. This
fourth year tuber produced 5-9 basal leaves (Fig. 2
E-F and Fig. 3) and their size varied from 10-12 cm.
The dissections of the leaves increased and the size
of the ultimate leaf segments were observed to be
variable. During fourth year of growing season, the
tuber turned darker in color and attained diameter
up to 3.5 cm. The size of the fourth year tuber was
comparable to the size of the tuber transplanted
from natural populations. During the same year
(4th years) these tubers initiated shoot development,
which subsequently developed flowers in the orders
starting from primary to higher orders.
3.2. Seed morphology
Like all umbellifers, the species studied now,
is characteristic in having a schizocarpic fruit; the
seed coat is fused with the fruit wall. The fruit is the
dispersal unit, which on maturity splits into two
European Journal of Biological Research 2016; 6 (3): 145-151
148 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
mericarps. Each mericarp alongwith the seed forms
a single dispersal unit. The fruit is subdidymous,
broad at base and narrow towards apex where it is
capped by a stlypodium from which two recurved
styles emerge. Each mericarp carries five primary
ridges, one dorsal, two commissural and two
intermediate. The ridges are corky, obtuse and run
parallel to the longitudinal axis of the mericarp. In
between the primary ridges, are present vallecular
spaces.
Seed weight: The seeds are small in size; each
mericarp measures 1.5-2.3 (x̄ 2.1) mm in length and
0.8-1.1 (x̄ 0.09) in breadth, ( commissural face). The
seed weight of 100 mericarps varies between 70-95
(x̄ 90) mg.
percent in Tikri and x̄ 60.2 ± 0.51 (0-65.0) in Reasi
population. The fruit set was also recorded separately for normal disomics and those bearing 1-4
extra chromosomes, beside the norma complement
(Table 1).
The data given in Table 18 reveals that in
comparison to normal disomics, the percentage fruit
set reduces, with increase in the number of extra
chromosomes, in all the umbel orders. In primary
umbel it reduced from 93.47 in disomics to 52.13
percent in plants with 2n=20 ± 4 extra chromosomes. Similar trend was observed in the fruit set of
secondary, tertiary and quarternary umbel orders.
3.3. Seed set
As the seed germinate, the radical is the first
to emerge and grow geotropically. Subsequently,
the two cotyledons emerge and on reaching above
ground (Fig. 2 A-B) start turning green. The cotyledons are unequal sized in majority of the seedlings
and remain green for 20-30 days. During this period
the seedlings attain a length of 5.0-5.4 cm. The
primary root also grows in size, which varies from
3.0-3.4 cm. As the elongation of primary root is
taking place; the development of a tuber is initiated
in the form of a small swelling at its sub-apical
position (Fig. 2 C). However, there is no shoot
formation. Towards the end of the growing season,
the cotyledons wither and the seedlings perennate
through the tuber. The epidermis of the tuber turns
light brown. As the tuber sprouts during next
growing period, a basal leaf emerges from the tuber
and reaches above ground (Fig. 2 D).
Although all seedlings at the stage carried a
single basal leaf but the dissection of the leaf was
variable. The basal leaf of these second seedlings
remained green for 30-40 days. During this period,
photosynthesis in leaves provided nutrition to
developing tubers, which attained a size of 8 to 9.5
mm. The leaf withered towards end of the growing
period and the tuber perennated in the soil, upto 15
cm deep from the ground level.
During third year growing season, the tuber
again sprouted and formed 2-3 basal leaves (Fig. 2
E-F). After further growth, the basal leaf emerges
above the soil and turned green. The mature leaves
formed during third year were more dissected and
large in size (9-10 mm).
Scaligeria stewartiana (Nasir) Nasir exhibits
variation in seed set at intra- as well as at inter
population level. The umbels of different order
portray variation in their contribution to the total
fruit yield of an individual plant. In Mansar
population, the tertiary umbels contribute maximum
to the total fruit yield of the individual plant. On
the contrary secondary umbels are the major
contributors to the fruit yield per plant in Tikri and
Reasi populations. However, irrespective of the
contribution of different umbel orders to the total
fruit yield, the fruit set decreases from primary to
the higher order umbels in all the three population.
The average fruit set in primary umbel of all
the three populations was above 90 percent. It was
x̄ 90.62 ± 0.55 (77.7-100) percent in Mansar population, x̄ 93.95 ± 0.48 (75.0-100) percent in Tikri
population and x̄ 93.38 ± 0.26 (80.0-100) percent
population. Percentage fruit set in secondary order
umbels varied between 88-90 percent, being x̄ 88.9
± 0.35 (72.0-100) in Mansar, x̄ 88.48 ± 0.32 (81.895.0) in Tikri, x̄ 90.94 ± 0.23(79.0-100) percent in
Reasi population.
The fruit set in third order umbels ranged
between 70-83.38 percent. The Mansar population
had 70.31 ± 1.49 (49.6-100) percent whereas in Tikri
and Reasi populations, fruit set was x̄ 83.38 ± 0.71
(70-100) and x̄ 81.24 ± 0.36 (36.2-90) percent
respectively. There was wide variation in the fruit
set of fourth order umbels. It was x̄ 65.72 ± 1.36
(0-90.6) percent in Mansar, x̄ 78.1 ± 0.49 (0-88.8)
3.4. Seedling establishment
European Journal of Biological Research 2016; 6 (3): 145-151
149 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
Table 1. Fruit set at umbel in Scaligeria stewartiana (Nasir) Nasir.
Primary order
Chromosome
number (2n)
20
20+1
20+2
20+3
20+4
Number
Percent
fruit set
Bisexual Fruit
Flower formed
x̄ 160±
149±
93.47±
1.31*
1.21
0.24
(110(91(82.6179)**
179)
100)
159±
129±
81.43±
6.6
5.35
1.91
(136(104(76.5163)
144)
88.8)
138±
87±
63.25±
12.4
7.81
5.90
(126(90(72-81)
225)
182)
132±
80±
60.1±
9.6
6.5
4.91
(125(65(52-70)
169)
118)
130±
67±
52.13±
8.9
4.2
3.24
(120(54(45-80)
175)
140)
Secondary order
Number
Bisexual Fruit
Flower formed
340±
326±
9.25
4.27
(170(152660)
660)
370±
287±
8.93
6.91
(175(121710)
642)
410±
195±
10.68
5.08
(190(81690)
340)
390±
180±
11.62
5.36
(180(72560)
276)
480±
227±
11.58
5.47
(225(86720)
175)
Tertiary order
Percent
fruit set
95.94±
0.28
(89.9100)
77.6±
0.83
(7090.47)
47.8±
0.54
(42.850.0)
46.2±
0.53
(40.149.2)
47.3±
0.63
(38.249.4)
Number
Quarternary order
Number
Percent
Bisexual Fruit fruit set Bisexual Fruit
Flower formed
Flower formed
396±
180±
72.23±
179±
127±
5.95
3.21
2.34
3.92
2.39
(250(120(48(0(0-275)
450)
490)
100)
253)
352±
126±
35.8±
171±
60±
7.43
2.46
2.0
4.23
2.47
(240(53(22.2(0(0-269)
560)
350)
62.5)
183)
360±
274±
45.15±
79±
29±
6.38
2.87
2.50
2.31
0.85
(255(117(28.6(0(0-182)
610)
360)
50)
124)
315±
77±
24.63±
69±
24±
5.27
4.21
9.6
2.11
0.74
(240(42.0(17.6(0-169) (0-61)
560)
280)
50)
290±
65±
22.59±
82±
28±
4.23
0.95
4.24
2.6
0.9
(210(32(15.2(0-159) (0-84)
510)
229)
45)
Percent
fruit set
71.2±
1.32
(4092.3)
35.0±
2.0
(067.9)
37.2±
2.3
(068.1)
35.2±
3.1
(069.5)
34.0±
3.43
(052.9)
Table 3. Dry biomass of Scaligeria stewartiana (Nasir) Nasir above and below ground parts of the seedings at different
stages of development.
Total dry
Dry biomass of below Dry biomass of above
Seedings
(N)
biomass (mg)
ground parts (mg)
ground parts (mg)
Ratio b:c
stage
a
b
c
x̄ 0.2330±0.0005*
0.1156 ± 0.0003
0.1174±0.0005
Ist year
20
1.01
(0.2021-0.2497)**
(0.1039-0.1251)
(0.1094-0.1279)
0.2286±0.0006
0.1137±0.0005
0.1149±0.0007
IInd year
20
1.01
(0.1935-0.2481)
(0.1120-0.1296)
(0.1012-0.1256)
0.1353±0.0003
0.1190±0.0005
0.1163±0.0006
IIIrd year
20
0.97
(0.1151-0.1532)
(0.1036-0.1295)
(0.1020-0.1347)
0.2570±0.0005
0.1230±0.0006
0.1110±0.0005
IV year
10
0.90
(0.2193-0.2695)
(0.936-0.1435)
(0.996-0.1239)
* Mean, S.E ** Range
These remained green and above ground for
35-40 days. During this period, the tuber also grew
in size upto 1.5 cm in diameter. During this period,
tuber also developed adventitious roots. The tuber,
which was almost globular during first and second
year, assumed irregular shape in the third year.
Towards the end of the growing season, the leaves
withered and tubers again entered into dormancy.
After perennation of the third year, the tubers
sprouted again during the growing season of fourth
year. The new basal leaves emerged above ground.
Each tuber produced 5-9 basal leaves (Fig. 2 A-B),
which varied 10.2-12.5 cm in length. The dissections of the leaves increased and the size of ultimate
leaf segments were also variable. The tuber became
darker in colour and attained diameter upto 3.5 cm.
The number of adventitious roots emerging from
each tuber also increased. The size of the fourth year
tuber was comparable to the size of tubers
transplanted from natural populations. During the
European Journal of Biological Research 2016; 6 (3): 145-151
150 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
same year (4th year) these tuber initiated shoot
development, which subsequently produced flowers
in the same order as was observed in case of tubers
transplanted from natural populations.
3.5. Seed viability
The viability of the seeds was determined by
tetrazolium test (Tz-test). The seeds of various
populations exhibited variation in their percentage
viability. In all the populations, seed viability was
above 85 percent (Table 4).
The strategy of resource allocation in the
species has been assessed after analyzing dry
biomass at various stages of seedling establishment
and seed set. The different stages of seedling exhibit
distinct morphology. In the subsequent years,
seedlings produce shoots and umbels of different
orders. The dry biomass of the seedlings at different
stages of development is represented in the Table 3.
Table 4. Seed viability in Scaligeria stewartiana (Nasir)
Nasir.
Seeds
Seeds
Population
Percentage
tested
viable
Botanic
95
89
93
Garden
Mansard
100
91
91
Tikri
120
102
85
Reasi
90
85
94.4
4. DISCUSSION
The life history pattern of a species is the
result of evolutionary decision in order to achieve
successful establishment, adaptation and future
evolutionary diversification [6]. The perpetuation of
a species depends at the rate at which individuals of
population contribute to future generations, which in
turn is influenced by the number of reproductive
attempts during its life time, age of first reproduction and reproductive rate. The herbaceous perennial
Scaligeria stewartiana (Nasir) Nasir produces
leaves and flowers every year from the underground
tuberous root which however it is not an organ of
vegetative multiplication [7]. The observations on
seed biology of the species have revealed that the
seed undergoes germination after attaining dormancy for the period of one year. After germination
of seeds the seedlings get established in four years,
after that every individual plant flowers for 4 to 5
years before perishing. The reproductive output also
increases with age, as the tuber increase in size. As
compared to only two umbel orders during first year
of reproductive age, the plant in the subsequent
years produce umbels upto fourth order [8]. During
first two years of seedling establishment, the species
invests more resources (dry biomass) in the formation of aerial parts and in third and fourth year, the
investment is more in the production of tuber. Long
gestation period in seedling establishment enables
the species to reduce competition and maintain
genetic composition of the populations. The weather
conditions are main cause of mortality. However,
permanent snow cover can protect seedling from
frost and heaving injuries [9-11].
5. CONCLUSION
Figure 4. Scaligeria stewartiana (Nasir) Nasir:
representing comparison of seed viability at different
locations in Jammu and Kashmir.
The populations of the species are decreasing
due to its seed dormancy and other types of
environmental disturbances, natural and human
interferences. The seed to seed cycle of the species
takes four years for completion whereas the total life
span of the species ranges from 7-8 years. During
first two years of seedling establishment, the
species invest more resources (dry biomass) in the
formation of ariel parts and in 3rd and 4th year, the
investment is more in the production of tuber. The
gestation period reduces competition between
European Journal of Biological Research 2016; 6 (3): 145-151
151 | Kour et al. Seedling establishment in Scaligeria stewartiana from North-West Himalayas, India
established individual and fresh recruits. However,
the species needs special attention due to a number
of threats.
3.
Kour K, Hamal IA, Gupta SK. Chromosomal
variability in Scaligeria stewartiana (Nasir) E.
Nasir. J Cytol Genet. 1992; 27: 43-45.
4.
Willson MF. Plant reproductive ecology. John
Wiley and Sons, Inc., New York, 1983.
5.
Koul V. Resource allocation in relation to floral
structure and breeding system in some member of
commelinaceace. Ph.D. Thesis, University of
Jammu, 1998.
6.
Putz N, Sakkau I. Seedling establishment bud
movement and subterranean diversity of subterranean system in Apiaceae. Flora. 2002; 197: 385393.
7.
Constance L, Chuang T, Bell CR. Chromosome
numbers in Umbelliferae. Am J Bot. 1971; 58(6):
577-587.
8.
Mukerjee PK, Constance L. Umbliferae of India.
Oxford and IBH. Publishing, New Delhi, 1993.
9.
Bergsten U, Nilsson JE. Early seedling growth of
Pinus sylvestris (L.) after sowing with a mixure of
stand and orchard seed in dence spacings. Can J
Forest Res. 2001; 31(7): 1184-1194.
ACKOWLEDGEMENTS
Authors are grateful to Prof. A. K. Koul, Dean
Academics, BGSB University, Rajouri for helpful
suggestions and thankful to Principal Govt. College
for Women, Parade Jammu for laboratory facilities.
AUTHORS’ CONTRIBUTION
KK: Data collection, Interpretation, Writing, Discussion; BLB: Manuscript designing, Discussion;
IAH: Guidance, Interpretation. The final manuscript
has been read and approved by all authors.
TRANSPARENCY DECLARATION
The authors declare no conflicts of interest.
REFERENCES
1.
Sarukhan J. Demography problems in tropical
system. Bot Monogr. 1980; 15: 161-188.
2.
Hamal, I. A. Cytotaxonomy of the umbellifers of
Kashmir Himalayas. Ph. D. Thesis. Jammu
University, 1981.
10. Bergsten U, Nilsson JE. Effects of seed weight and
seed type on early seedling growth of Pinus
sylvestris under harsh and optimal conditions. Scand
J Forest Res. 2002; 17: 118-130.
11. Vernet P, Harper JL. The costs of sex in seaweeds.
Biol J Linn Soc. 1980; 13: 129-138.
European Journal of Biological Research 2016; 6 (3): 145-151