Germination of some multipurpose tree species in five ... · insolação, da temperatura, substrato...

Tropical Ecology 46(2):203–217, 2005 ISSN 0564–3295 © International Society for Tropical Ecology

Germination of some multipurpose tree species in five provenances in response to variation in light, temperature, substrate and water stress

NEERAJ KHERA1 & R.P. SINGH2

Department of Forestry, N.D. University of Agriculture and Technology, Kumarganj Faizabad

Abstract: Seeds of Acacia catechu, A. nilotica, Albizia lebbek, Dalbergia sissoo and Tectona grandis from five provenances of northern India were tested to evaluate the effect of light quality, temperature, substrate and water-stress on germination of seeds, and to assess the extent of intraspecific variation. Results indicated that germination of Acacia catechu, A. nilotica and D. sissoo was less dependent on light quality, whereas direct and far-red light enhanced germination in some provenances of T. grandis and Albizia lebbek, respectively. Seeds of all the five species exhibited variation regarding the range of optimum temperature required for maximum germination. Small-sized seeds showed higher germination in paper methods, whereas germination of medium-sized seeds was higher in sand and soil methods. T. grandis exhibited very poor results in paper methods. Some provenances of A. nilotica and Albizia lebbek were found to be tolerant to significantly high levels of water stress. Intraspecific variation in A. lebbek, for temperature requirement and stress tolerance, was conspicuous.

Resumen: Semillas de Acacia catechu, Acacia nilotica, Albizia lebbek, Dalbergia sissoo y Tectona grandis de cinco procedencias del norte de la India fueron probadas con el fin de evaluar el efecto de la calidad de la luz, la temperatura, el sustrato y el estrés hídrico sobre la geminación de las semillas, y para evaluar la magnitud de la variación interespecífica. Los resultados indicaron que la germinación de A. catechu, A. nilotica y D. sissoo fue menos dependiente de la calidad de la luz, mientras que la luz directa y el rojo lejano promovieron la germinación en algunas procedencias de T. grandis y A. lebbek, respectivamente. Las semillas de las cinco especies mostraron variación en términos del intervalo óptimo de temperatura requerido para obtener la máxima germinación. Las semillas de tamaño pequeño mostraron una germinación más alta en métodos que usan papel, mientras que la germinación de semillas de tamaño mediano fue mayor en métodos que usan arena y suelo. T. grandis mostró resultados muy pobres en los métodos que usan papel. Se encontró que algunas procedencias de A. nilotica y Albizia lebbek son tolerantes a niveles significativamente altos de estrés hídrico. Fue notable la variación intraespecífica en A. lebbek relacionada con los requirimientos térmicos y la tolerancia al estrés.

Resumo: Sementes de Acacia catechu, Acacia nilotica, Albizia lebbek, Dalbergia sisso e Tectona grandis de cinco proveniências do Norte da Índia foram testadas para avaliar o efeito da qualidade da insolação, da temperatura, substrato e stress hídrico na germinação, e avaliar a extensão da variação inter-específica. Os resultados indicam que a germinação da A. cachu, A. nilotica e D. sissoo era menos dependente da qualidade da insolação enquanto que a insolação directa e a vermelha longa aumentou a germinação em algumas proveniências da T. grandis e A. lebbek, respectivamente. As sementes de todas as cinco espécies apresentaram variação em relação ao intervalo da temperatura óptima necessária para a germinação máxima. As sementes de pequenas dimensões mostraram mais elevada germinação para os métodos no papel, enquanto a germinação das sementes de média dimensão foram mais elevadas para os métodos em areia e terra. A T. grandis exibiu resultados muito pobres nos métodos em papel. Algumas proveniências da A. nilotica e Albizia lebbek apresentaram-se tolerantes a elevados valores de stress hídrico. A variação intra-específica na A. lebbek, para as necessidades de temperatura e tolerância ao stress foi conspícua.

Key words: Germination, intraspecific-variation, light, provenance, substrate, temperature,

waterstress.

Introduction

Corresponding Author: e-mail; [email protected] Present Address: 1Department of Natural Resources, TERI School of Advanced Studies, Darbasi Seth Block, India

Habitat Center, Lodhi Road, New Delhi 110003. 2Department of Forestry, Kumaun University, Nainital

INTRASPECIFIC VARIATION IN PROVENANCES 204

Introduction

Acacia catechu Willd., Acacia nilotica Willd., Albizzia lebbek Benth., Dalbergia sissoo Roxb. and Tectona grandis Linn., are some of the most extensively distributed tree species throughout the northern plains of India. These species are of special importance in the arid-dry regions of northern India. These are the most sought after species when it comes to afforestation of wastelands, avenue plantation or agroforestry purposes. Being cost effective, the commonly adapted technique for regeneration of these species is direct seeding. As, seed characteristics of a species with wide distribution may vary from place to place, the seed germination of different provenances is of great importance for natural and artificial regeneration. Further, the intraspecific variation in response of seeds to light, temperature, substrate requirements and stress tolerance can affect the afforestation programs significantly.

The germination of seed is strongly influenced by variation in temperature, water stress and light requirements, and these factors often show significant interaction in their effects on germination (Bokhari et al. 1975; Erasmus & Staden 1986). Media (germination substrate) also play a significant role in germination because seeds have characteristic requirements as to the amount of moisture and oxygen needed for germination (Purohit et al. 1998.).

A large number of studies have been carried out on the effect of quality of light, temperature regime, substrate and water stress on germination of forest tree species including the species covered in the present study. However, in none of the studies, intraspecific variation in germination requirement and stress tolerance among provenances has been studied. The present study analyses the intraspecific variation in different provenances of these species when exposed to different environmental conditions.

Materials and methods

Seeds were collected from five provenances of Acaica catechu, A. nilotica, Albizia lebbek, D. sissoo and T. grandis in northern India. These provenances were: Bahraich (P1), Gorakhpur (P2), north Gonda (P3), south Gonda (P4) and Faizabad (P5). Details of the provenances are given in Table 1. For each species, twenty healthy and apparently disease free trees, at least 100 m apart were selected in each provenance, and pods/fruits were collected during the time of maximum seed availability. After collection, seeds

were cleaned, extracted and made homogenous to form five lots representing five provenances of that particular species.

Germination studies were conducted at the main Experiment Station, N. D. University of Agriculture & Technology, Kumarganj, Faizabad, in Northern India, which is situated at an elevation of 113 m above mean sea level at 260 47’

N lat and 820 12’ E long. The climate is tropical with an average annual rainfall of 1022 mm.

In all experiments, a factorial design with four 25-seed replicates was used. For germination, seeds were placed on cut sheets of germination paper in Petridishes and moistened with 5 ml of distilled water (ISTA 1985).

Light quality Red light was obtained by wrapping the Petri

dishes with red cellophane paper, while wrapping the Petri dishes with blue and red cellophane papers created far-red light conditions. Dark conditions were obtained by wrapping Petri dishes with two black carbon papers. Petri dishes covered with transparent cellophane sheets were used as control. 100 W incandescent bulb kept at a distance of approximately 1 m provided illumination (Rao 1986). These experiments were carried out in a dark room and low intensites of green light were used during handling. These tests were conducted at laboratory temperature.

Temperature The effect of temperature on the germination

of seeds was studied under following temperature regimes: 200C, 250C, 300C, 350C and 400C constant and 200/300C and 250/350C fluctuating tempera-tures, with 8 hours of higher range and 16 hours of lower range. The Petridishes were kept in the germination chamber with 8 hours of photoperiod. In case of fluctuating temperatures, photo-period was kept during the higher range of temperature.

Germination substrate Seeds were set for germination in six different

substrates. These substrates were: a) Top of the Paper (TP), where seeds were placed over the sheet of germination paper; b) Between Paper (BP), where seeds were placed between two sheets of germination paper; c) Top of Sand (TS), where sand was filled in enamel tray and seeds were placed over sand. After placing seeds, a thin layer of sand was placed covering the seeds completely; e) Top of Soil (TSI), same as TS method; and f) Soil (SI), same as sand method. Among these a and b were set in Petridishes and

KHERA AND SINGH

205

( )

Table. 1. Location of the provenances studied. Provenances Location Altitude Rainfall Temperatures (0C)

(m) (mm) Maximum Minimum P1 Bahraich 27030’N 81031’E 124 1177 37.5 9.8 P2 Gorakhpur 27054’N 82024’E 117 1044 34.8 10.1 P3 North Gonda 27024’N 82005’E 119 1330 37.0 10.5 P4 South Gonda 27024’N 82005’E 119 1245 37.0 10.5 P5 Faizabad 26046’N 82023’E 112 1142 39.0 8.4

c-f were set in enamel trays. In order to prevent the variation in the moisture level of substrata during the test period, all the enamel trays were covered with transparent sheets of cellophane paper and placed in the germination chambers along with the Petri dishes.

Seeds were set for germination in Petridishes as in the experiments of light and temperature. But the germination papers were moistened with different concentrations of mannitol solution to maintain water poetential of -5, -10, 15 and -20 bars (Uhvits 1946). Various levels of water stess were made according to the formula given by Helmerick & Pfeifer (1954). Solution of mannitol of respective concentration was added to the Petri-dishes whenever required during the experiment. Petri dishes where distilled water was used served as control.

In all the experiments, observation on seed germination were made after every three days up to 90 days in case of T. grandis and 60 days for the rest of the species. A seed was considered as germinated when the radicle had protruded about 2 mm beyond the seed coat. In the experiments involving sand and soil as substrate, emergence of radicle just at the surface was considered as germination. Germination value (GV) was calculated using Czabato (1962). Germination Energy (GE) was expressed as percent germination after 6 days in A. catechu, A. nilotica, A. lebbek and after 60 days in T. grandis (Ford-Robertson 1971). Measure of response breadth was calculated using Levin’s niche breadth metric (Levin 1968) as:

Water stress

sPsB

i 1

2∑=

i=1 where, Pi is the proportion of seed

germination response in state i, and S is the total number of states (treatments). The resulting measure B is a scale from 0 to 1 being the widest breadth.

Results and discussion

Light Results indicate that all the five provenances

of A. catechu exhibited a common trend in which the values of percent germination, GV and GE were highest under direct light condition (Table 2). However, germination was equally good under far-red light in P1, P2 and P3. Germination was significantly reduced in dark conditions in all provenances except P4. Provenance P5 differed from other provenances as germination was significantly reduced in red, far-red and dark conditions.

Out of five provenances of Acacia nilotica, three provenances exhibited similar trend, as there was no significant variation in germination under different light conditions. These three provenances are P1, P3 and P4. However, in P2 significantly higher germination was observed under far-red light and significantly reduced germination in dark. On the other hand, P5 exhibited significantly reduced germination in dark conditions.

Provenances of A. lebbek differed in their germination response to different light conditions. P1, P2 and P3 exhibited better germination in far-red light whereas, in P4 and P5, it was higher in red light. However, in all the five provenances, germination was greatly reduced in dark.

Seeds of D. sissoo from all the five provenances exhibited a significantly reduced germination under dark conditions. However, the provenances differed in the light quality in which they exhibited higher germination. In P1, P2 and P5, there was no significant variation in germination under direct, red and far-red light; in P3, germination was equally higher in direct and far-red light and it was greatly reduced in red light; in P4, germination was significantly different in all the light conditions. In T. grandis, germination was significantly higher under direct light as compared to other light conditions. (Fig.1).

Tab

le 2

. Ger

min

atio

n of

see

ds fr

om d

iffer

ent p

rove

nanc

es u

nder

diff

eren

t lig

ht c

ondi

tions

. Pr

oven

ance

Ligh

tAc

acia

cat

echu

Ac

acia

nilo

tica

Albi

zia

lebb

ekD

albe

rgia

sis

soo

Tect

ona

gran

dis

G

%

GV

G

E (%

) G

%

GV

G

E (%

) G

%

GV

GE

(%)

G%

G

V

GE

(%)

G%

G

V

GE

(%)

P1

Dir

ect

93

218.

2 93

91

19

.02

5466

11.5

37

9455

9.4

94

140.

014

9

206 INTRASPECIFIC VARIATION IN PROVENANCES

Red

85

163.

9 85

90

10

.26

53

81

16.9

4796

521.

296

90.

012

9Fa

r-re

d90

20

1.5

90

95

16.8

4 45

90

48.4

4796

714.

596

40.

001

3D

ark

87

19

7.3

78

90

18.2

9 50

49

6.07

3385

523.

485

70.

001

6

P2D

irec

t89

19

9.6

89

86

33.0

2 63

3214

.620

2816

.128

220.

391

19R

ed67

15

2.5

67

88

44.8

0 72

24

10

.019

2415

.324

120.

036

12Fa

r-re

d85

17

9.5

85

94

21.7

5 84

30

8.33

2022

12.3

2210

0.02

510

Dar

k

79

183.

4 79

69

11

.05

38

82.

678

189.

8210

90.

020

9

P3D

irec

t97

22

4.5

97

94

136.

24

8473

29.4

4463

85.2

6312

0.01

98

Red

82

167.

8 82

94

63

.15

85

42

5.90

2254

74.5

548

0.00

85

Far-

red

92

191.

5 92

92

66

.90

75

7629

.237

6180

.461

100.

012

8D

ark

89

20

1.7

89

91

120.

21

76

589.

5020

5056

.216

40.

002

4

P4D

irec

t95

23

1.6

95

94

69.2

3 77

435.

4226

7832

5.0

7818

0.26

010

Red

81

152.

1 75

91

60

.45

82

58

6.45

3265

266.

965

150.

189

9Fa

r-re

d89

18

4.3

89

91

89.9

6 78

46

4.20

2670

196.

470

110.

093

5D

ark

92

21

0.4

92

90

55.7

6 65

34

4.34

2154

98.3

054

100.

084

3

P5D

irec

t94

22

5.6

94

8218

.4

7340

3.84

2184

252.

584

60.

002

6R

ed66

14

3.6

58

73

11.6

5 48

51

7.

0030

8325

0.3

834

0.00

22

Far-

red

84

156.

2 85

80

33

.77

75

373.

9221

7923

3.6

793

0.00

11

Dar

k69

15

8.2

61

79

11.8

7 49

28

1.98

1576

231.

776

30.

001

2

CD

for

A 5.

2332

.93

5.23

5.94

8.9

9.07

5.78

3.2

6.83

6.52

55.3

46.

52

4.49

0.04

4.26

C

D fo

r B

5.

3833

.81

5.38

6.09

9.15

7.17

5.96

3.27

7.04

6.68

56.8

16.

68

4.62

0.04

4.39

C

D fo

r A

x B

8.

2451

.88.

249.

3114

.05

11.0

99.

095.

0410

.79

10.2

787

.11

10.2

77.

100.

066.

67 A

= P

rove

nanc

e, B

= L

ight

qua

lity;

GV

= g

erm

inat

ion

valu

e; G

= ge

rmin

atio

n

KHERA AND SINGH 207

Fig.1. Germination behavior of seeds under different light conditions (data averaged across provenances)

Response breath were wider for light condition in A. catechu, A. nilotica and D. sissoo, and comparatively narrow in A. lebbek and T. grandis, showing that the germination of the former three species is less dependent upon the type of light quality (Table 3).

Temperature All the five species exhibited a common trend

in which germination was significantly low at 200C (Fig.2). It increased from 200C to higher temperatures, reached a peak point and then declined at further high temperatures. This may be due to the fact that as the temperatures rise, changes in protein conformation occur which actually promote the germination process, but further conformational changes occur which are deleterious to it as temperatures become too high (Bewley & Black 1985).

Provenances differed in the peak point, i.e. the temperature for highest germination. In A. catechu, seeds from P1, P2, P3 and P5 exhibited highest germination in the constant temperature range of 30-350C and fluctuating temperature of 20/300C. However, temperature range for the seeds of P4 was 30-400C constant and fluctuating temperature of 20/300C. Germination of seeds of A. nilotica was not affected significantly by temperature of 20/300C. Germination of seeds of A. nilotica was not affected significantly by

temperature variation in the range of 25-400C, it was highest at 25/350C fluctuating temperature. Among the five provenances of Acacia nilotica, P4 emerged as the most tolerant one, as the rise or decline in germination over different temperatures was not significant. Tolerance to a wide range of temperature may help the species to germinate in adverse climatic conditions and extend its natural distribution range. Kozlowski (1970) and Osmond et al. (1980) also stated that species, which germinate readily over a relatively wide range of temperatures, should be easier to establish in the field than those with highly specific temperatures.

A temperature range of 30-350C was found to be favourable for seed germination of D. sissoo in all the provenances.

Different provenances of A. lebbek and T. grandis exhibited differential germination response to various temperature ranges (Table 4). In case of Albizia lebbek, germination was highest at 20/300C and 25/350C fluctuating temperatures in P1, whereas P2 exhibited highest germination at 25/350C. In P3 and P4, germination was higher at 30-350C constant temperature and 25/350C fluctuating temperatures. In P5, the germination was highest at 350C and 25/350C. It shows that P1 had a wider tolerance range for temperature. In T. grandis seeds of P2 were found to have wider tolerance range for temperature.


It breathniloticrangetempefairly The dnilotictemperisingconne1973) rise inincreagermiand tempethose

Table 3. Levin’s B (response breadth) for different species for various parameters on differentenvironmental factors.

Species Provenance Environmental factors Light Temperature Substrate Water Stress A. catechu P1 1.00 0.98 1.00 0.98 P2 0.99 0.99 0.99 0.99 P3 1.00 0.99 0.99 0.99 P4 0.99 1.00 1.00 1.00 P5 0.98 0.99 0.99 0.99 Average 0.992 0.990 0.994 0.990 A. nilotica P1 1.00 1.00 0.99 0.99 P2 0.99 1.00 0.98 0.99 P3 1.00 1.00 0.98 1.00 P4 1.00 0.99 0.99 1.00 P5 1.00 1.00 0.99 0.99 Average 0.998 0.998 0.986 0.994 A. lebbek P1 0.96 1.00 0.99 0.90 P2 0.85 0.88 0.85 0.98 P3 0.96 0.83 0.96 0.98 P4 0.97 0.82 0.95 0.98 P5 0.96 0.83 0.94 0.96 Average 0.940 0.872 0.938 0.960 D. sissoo P1 1.00 1.00 0.81 0.95 P2 0.98 0.99 0.90 0.86 P3 0.99 0.99 0.93 0.98 P4 0.98 0.99 0.78 0.84 P5 1.00 0.99 0.92 0.81 Average 0.990 0.992 0.868 0.888 T. grandis P1 0.85 0.96 0.73 0.69 P2 0.87 0.87 0.78 0.60 P3 0.91 0.89 0.71 0.62 P4 0.95 0.89 0.72 0.63 P5 0.92 0.79 0.76 0.55 Average 0.900 0.880 0.740 0.618

is evident from the expression of response s that among the five species, seeds of A. a showed maximum tolerance to a wide of temperatures as, except very low ratures, germination of this species was good at all temperatures in all provenances. ecline in the germination percentage of A. a and T. grandis after a certain rature, when germination value was still , may be due to the fact that a causal ction may exist (Gulliver & Heydecker between the decline in percentage and the rate, as high temperatures are likely to

se the rate of many component processes of nation. In seeds in which they still function are well coordinated at the higher ratures the rate of germination increase, in which they are not will tend to fail.

Except D. Sissoo and A. catechu, all the species exhibited inverse relationship between temperature and occurrence of germination. Bernstein (1975) and Rana (1977) have emphasized the importance of early initiation and completion of germination process since they found that this phase is relatively more sensitive and often more decisive than subsequent growth stages.

Germination substrate Germination substrate was found to play a

significant role in the germination behaviour of seeds of all the five species (Fig.3). Seeds of A. catechu from all provenances, except P5, exhibited higher germination in BP, TP and TS methods. In P5 germination was not affected by the type of substrate (Table 5). Seed germination

KHERA AND SINGH

209

in P1 and P5 provenances of A. nilotica did not vary in different substrates. However, in P2 and P3, germination was significantly low in sand and soil methods. P4 exhibited slightly less germination in soil method.

Germination was quicker in paper method in all species excepting T. grandis, in which germination was quicker in soil method (Table 5). This may be due to the fact that seed of this species being large and round shaped were not properly moistened in paper methods. Proper moistening of teak seeds is perhaps an essential requirement to germination (Kehera, 2004 unpublished)

Seeds of A. lebbek from all provenances exhibited significantly high germination in BP and TS methods. P4 showed higher germination in BP and sand method. Seeds of D. sissoo exhibited higher germination in paper methods, however, P1 and P5 showed highest percent germination and GV in BP, whereas P2, P3 and P4 showed their highest values in TP method.

In case of T. grandis, the effect of substrate was more pronounced as compared to other species. Seeds from all the provenances exhibited higher germination in soil method, and

significantly reduced germination in paper methods.

Fig. 2. Germination behaviour of seeds under different temperature (data averaged across provenances).

Response breaths were wider for A. catechu and A. nilotica, showing their adaptability to various types of germination substrates; slightly narrow for A. lebbek and D. sissoo indicating that germination of these species is influenced by the type of germination substrate (Table 3). T. grandis exhibited a narrow (0.72) response breath depicting that germination of this species is largely dependent on the type of germination substrate. It was also observed in the study that, species with smaller seed size, viz., A. catechu and D. sissoo exhibited higher germination in paper methods as compared to sand and soil methods. Germination percentage, in both species, was fairly high in TS method also but germination value was remarkably less. This may be due to the mechanical obstruction posed by the sand particles on the emerging radicle, resulting in lower values of GV and GE. Also, seeds of D. sissoo have very delicate and thin seed coat, which is likely to get damaged by sand abrasion, resulting in poor germination percentage. Aldhous (1972) also found that sand

is not suitable for very small seeds. Higher germination in paper methods for small sized seeds have also been reported for Alnus neplaensis. On the other hand, A. nilotica and A. lebbek performed fairly good in both paper and TS methods. This may be due to the fact that these species have thick seed coat and large sized seeds, which require proper moisture conditions in order to get rid of seed-coat dormancy. Further as the germination periods is also longer in these species, sand and soil restrict the spread of fungi. Catalan (1992) also found that in Prosopis flexuosa and P. chilensis the use of either sand or paper gave similar results.

s not suitable for very small seeds. Higher germination in paper methods for small sized seeds have also been reported for Alnus neplaensis. On the other hand, A. nilotica and A. lebbek performed fairly good in both paper and TS methods. This may be due to the fact that these species have thick seed coat and large sized seeds, which require proper moisture conditions in order to get rid of seed-coat dormancy. Further as the germination periods is also longer in these species, sand and soil restrict the spread of fungi. Catalan (1992) also found that in Prosopis flexuosa and P. chilensis the use of either sand or paper gave similar results.


T. grandis exhibited very poor germination when paper was used as substrate, while germination was high using soil and sand as substrates. This was in contrast to the findings of Purohit et al. (1998) who reported that large as well as medium or small sized seeds showed better germination in both Paper and TS method. The reason of poor germination of teak in paper method is that teak seeds are quite large sized, and moreover they need sufficient humidity to germinate (Khera 2003, unpublished), and it is not possible to cover the surface of teak seeds completely in paper methods.

T. grandis exhibited very poor germination when paper was used as substrate, while germination was high using soil and sand as substrates. This was in contrast to the findings of Purohit et al. (1998) who reported that large as well as medium or small sized seeds showed better germination in both Paper and TS method. The reason of poor germination of teak in paper method is that teak seeds are quite large sized, and moreover they need sufficient humidity to germinate (Khera 2003, unpublished), and it is not possible to cover the surface of teak seeds completely in paper methods.

Fig. 3. Germination behavour of seeds under different substrates (data averaged across provenances).

KHERA & SINGH 211

Tab

le 4

. Ger

min

atio

n of

see

ds fr

om d

iffer

ent p

rove

nanc

es in

diff

eren

t tem

pera

ture

s.

Prov

enan

ceTe

mpe

ratu

re

Acac

ia c

atec

hu

Acac

ia n

ilotic

aAl

bizi

a le

bbek

Dal

berg

ia s

isso

oTe

cton

a gr

andi

s

G%

G

V

GE

(%)

G%

G

V

GE

(%)

G%

G

V G

E (%

) G

%

GV

G

E (%

) G

%

GV

G

E

(%)

P1

20

0 C85

81.1

069

853.

2112

321.

910

8115

1514

0.02

1 6

250 C

85

214.

2 85

89

14.6

3 26

41

9.2

2180

1414

9

0.06

3 10

300 C

94

221.

0 94

90

29.4

6 57

60

19.5

37

9421

214

0.34

6 17

350 C

94

226.

1 94

90

100.

6 63

68

49.5

39

9324

247

0.89

4 24

400 C

87

172.

4 87

91

199.

6 84

67

64.6

50

8423

23

0.90

1 23

200 /3

00C

9619

1.4

9689

183.

469

7076

.349

9015

150.

321

1525

0 /350

C

9720

5.9

97

9620

4.3

9074

114.

160

8124

240.

462

24

P220

0 C78

92

.10

78

761.

76

122.

14

204.

906

90.

016

625

0 C77

17

9.9

77

7819

.3

28

199.

68

2211

.422

160.

084

1230

0 C88

19

6.4

88

8063

.0

73

2320

.321

2313

.123

300.

541

2835

0 C89

20

4.2

89

8158

.4

64

2935

.621

2312

.923

350.

726

3340

0 C81

19

4.3

81

7772

.9

41

1619

.512

189.

6018

430.

929

4020

0 /300

C83

199.

883

7861

.261

3031

.429

1910

.419

260.

304

2625

0 /350

C

6715

3.1

6782

81.3

7641

61.2

3415

8.30

1529

0.49

028

P320

0 C89

54.3

0 68

83

4.6

12

211.

95

5740

.115

50.

003

225

0 C89

20

9.4

89

8471

.2

32

346.

912

6075

.060

110.

015

930

0 C98

22

6.2

98

9110

1.2

81

7336

.248

6588

.465

180.

391

1635

0 C97

23

0.4

97

8798

.6

80

7659

.454

6487

.264

240.

529

2240

0 C84

19

1.6

84

8056

.5

78

2013

.518

5079

.650

210.

601

2020

0 /300

C98

216.

598

8810

6.9

7356

26.2

4355

83.6

5516

0.31

116

250 /3

50C

8810

0.5

8894

126.

385

7864

.468

4859

.948

200.

409

20

C

ontin

ued…

INTR

ASP

EC

IFIC

VA

RIA

TIO

N IN

PR

OV

EN

AN

CE

S 21

2

Tabl

e 4.

Con

tinue

d

P4

20

0 C79

93.4

0 51

88

6.1

388

1.0

372

214.

072

100.

031

10

250 C

85

221.

5 85

90

25.7

45

24

8.4

1280

361.

180

120.

094

1130

0 C96

23

6.1

96

9317

5.6

89

7217

.130

7935

1.1

7926

0.51

319

350 C

98

249.

4 98

95

268.

4 89

75

68.9

6175

312.

475

320.

634

3240

0 C97

22

0.1

97

9643

7.3

96

4959

.940

6528

6.1

6530

0.69

930

200 /3

00C

9825

3.6

9894

306.

481

6143

.250

7630

2.1

7621

0.43

620

250 /3

50C

8920

1.4

89

9739

8.6

89

7464

.468

5922

5.6

5927

0.53

122

P520

0 C75

52.1

0 43

79

2.0

5 12

1.3

571

95.9

425

0.00

32

250 C

8422

0.2

84

8119

.4

13

312.

112

7819

8.3

784

0.00

32

300 C

9422

4.2

94

8089

.6

62

433.

625

8425

4.6

8414

0.39

014

350 C

9523

0.6

95

8698

.5

67

6424

.630

8325

0.4

8324

0.45

924

400 C

7618

0.4

76

8810

4.2

72

3514

.922

7018

6.2

7020

0.53

220

200 /3

00C

9423

6.1

9485

97.9

6530

14.2

1580

223.

180

130.

319

1325

0 /350

C

8120

3.4

8191

121.

485

6931

.433

6514

2.4

6520

0.40

120

C

D fo

r A

3.71

21.1

3.98

3.

1212

.64.

645.

085.

664.

535.

736

.83

5.66

4.22

0.08

3.91

C

D fo

r B

3.

8525

.01

4.16

3.27

13.1

44.

855.

296.

084.

755.

9528

.39

5.88

4.41

0.07

4.09

C

D fo

r A

x B

7.

3347

.68

7.95

6.22

25.0

59.

2410

.09

11.2

39.

0311

.35

73.1

411

.23

8.39

0.18

7.70

Tab

le 5

. Ger

min

atio

n of

see

ds o

f diff

eren

t pro

vena

nces

, on

diffe

rent

type

s of

ger

min

atio

n su

bstr

ate.

Prov

enan

ce

Tem

pera

ture

Ac

acia

cat

echu

Ac

acia

nilo

tica

Albi

zia

lebb

ekD

albe

rgia

sis

soo

Tect

ona

gran

dis

G

%

GE

(%)

GV

G

%

GV

GE

(%)

GV

G

%

GE

(%)

G%

G

V

GE

(%)

G%

G

V

GE

(%

) P1

B

P93

21

8.2

93

9029

.2

53

5919

.736

9458

2.8

945

0.00

2 5

TP92

18

8.5

92

90

18.9

49

67

10

.139

6620

1.3

662

0.00

12

TS92

15

9.1

92

96

40.1

43

73

19

.238

7591

.43

6314

0.01

38

S87

14

6.3

72

82

27.3

25

74

30

.552

181.

016

210.

342

17TS

I91

14

4.9

91

94

19.8

27

56

22.4

2876

80.4

2417

0.25

612

SI86

13

9.8

68

90

20.7

9 62

13

.828

230.

30

280.

521

28

Con

tinue

d…


KH

ER

A A

ND

SIN

GH

213

199.

6 83

81

.6

22

21

20

7

Tabl

e 5.

Con

tinue

d

P2B

P91

91

72

22

19.6

10.1

0.

002

7

18

6.2

85

31.3

13

.9

22

31

31

4

3

146.

4 78

39

.0

15.4

30

31

24

21

19

14

1.1

64

15.6

13

13

30

28

137.

6 82

17

.9

10.6

28

26

0

26

25

13

2.0

57

13

3.0

0 31

30

22

4.5

95

88.9

46

62

60

4

4

220.

5 94

14

9.6

30.2

47

74

74

1

1

19

3.6

92

52.6

25

.2

54

70

0 12

8

18

7.7

73

24.4

13

.7

36

43

32

18

16

18

9.4

86

37.3

10

.1

27

47

0 19

18

177.

9 68

10

.4

21

30

2 0

25

25

23

1.6

96

160.

0 31

80

80

3

3

22

2.4

94

67.9

82

82

2

1

20

5.2

88

135.

3 49

77

54

18

10

217.

0 94

10

1.8

17.5

38

31

19

25

19

201.

3 80

24

.9

63

4.9

0 24

16

197.

3 80

43

.6

45

2 3.

0 0

29

29

22

5.6

67

31.9

21

85

81

3

3

20

2.4

72

42

80

76

3

2

208.

1 71

49

80

65

6

6

197.

9 68

28

32

29

14

14

194.

5 74

25

58

29

.9

0 11

11

179.

7 72

24

0.

9 8

51

34.8

0

15

15

13

.17

7.80

10

.97

1.59

TP90

90

73

30

39

.60.

001

TS88

88

63

46

19

13

.50.

361

S87

69

44

3.

0 14

3.

3 0.

530

TSI

84

84

55

34

9.0

0.45

6

SI

81

52

5.

4 9

12

0.80

6

0.80

9

P3B

P97

97

86

72

35.0

84.8

0.

003

TP97

97

86

74

28

3.8

0.00

1TS

95

95

72

76

39.2

0.

019

S90

79

42

48

25

.5

0.38

4TS

I93

93

78

47

12

.4

0.40

5

SI

87

68

36

50

5.

9

0.53

9

P4B

P95

95

93

72

16

.9

40

324.

60.

002

TP95

95

91

5.

9 26

35

0.9

0.00

1TS

94

94

83

9.1

20

85.7

0.

253

S91

80

78

58

41

28

.6

0.50

1TS

I93

93

56

4.

6 20

0.

483

SI87

78

49

6.

5 26

0.

541

P5B

P94

94

38

41

3.1

254.

9 0.

002

TP94

94

16

.8

42

4.3

24

219.

30.

001

TS94

94

13

.2

26

7.0

23

86.2

0.

002

S92

78

12

.9

24

3.1

16

27.2

0.

393

TSI

91

91

15.0

28

1.

56

0.40

1

SI

89

65

15

.1

34

0.42

4

C

D fo

r A

3.39

26.9

55.

089.

267.

154.

264.

732.

925.

980.

651.

494.

61

4.06

0.

084.

10

CD

for

B

3.44

27.3

94.

929.

387.

254.

314.

822.

976.

050.

711.

634.

63

4.13

0.

074.

17

CD

for

A x

B

6.25

49

.56

9.36

17

.01

16.4

7 5.

37

3.65

7.

95

7.48

0.

15

7.57

KHERA & SINGH 213

Intraspecific variation was observed in A. nilotica, A. lebbek and D. sissoo, as seeds from different provenances differed in their germination response to different types of germination substrate.

Water stress In A. catechu, germination declined with the

increased water stress, however, all the five provenances differed in their germination response to different levels of water stress, P1 emerged as the most tolerant with a threshold

between -10 to -15 bars, whereas for all other provenances the threshold was between -5 to -10 bars (Table 6).

Fig. 4. Germination behavior of seeds in different substrates (data averaged across provenances)


Some provenances of A. nilotica and A. lebbek exhibited an increased germination at moderately higher levels of water stress. In A. nilotica, though GV declined with increase in stress, percent germination of P1 and P3 increased significantly with an increased stress up to -5 bars. In all other provenances, there was no significant decline in percent germination up to -5 bars. Even at higher levels of stress, all

ce

Wa

Tab

le 6

. Effe

ct o

f diff

eren

t lev

els

of w

ater

str

ess

on g

erm

inat

ion

beha

vior

of s

eeds

. Pr

oven

ante

r st

ress

(bar

) Ac

acia

cat

echu

Ac

acia

nilo

tica

Albi

zia

lebb

ek

Dal

berg

ia s

isso

o Te

cton

a gr

andi

s

G

%

GV

G

E (%

) G

%

GV

G

E (%

) G

%

GV

G

E

(%)

G%

G

V

GE

(%

) G

%

GV

G

E

(%)

P1

0 22

1.0

90

29.0

1 58

59

93

93

22

17

94

94

20.6

37

579.

4 0.

351

197.

4 23

.66

49.2

73

89

89

20

13

19

4.5

24.7

2 38

.9

68

64

52

14

150.

2 15

.09

17.6

53

60

19

6

4 97

.8

11.7

1 16

53

89

.5

8 0

P2

0 19

6.4

80

63.9

6 72

23

20

.3

33

21

11.2

16

30

28

159.

9 23

.40

25.1

18

20

10

.2

15

19

17

14

3.6

18.9

4 16

.3

17

15

9.5

7 11

11

142.

1 18

.19

11

6.2

6 5

5 94

.6

14.6

8 3

5 3.

2 2

0 0

0

P3

0 22

6.2

80

47

62

55

18

15

20

5.6

26.2

9 31

.3

59

70

62

15

19

4.2

28.2

9 30

.45

69

68

60

8

6

16

4.6

33.3

3 20

.5

61

57

80.6

40

3

2 98

.3

22.2

1 43

44

35

.9

31

0

P4

0 23

6.1

90

30

79

79

26

19

214.

5 95

.7

13.8

36

37

86

.2

12

20

16

197.

4 94

.8

10.0

34

32

76

.6

11

11

6

18

4.3

29.1

9 32

69

.0

12

5 2

103.

2 27

.10

23

31

17.2

9

0

P5

0 22

4.2

75

14.3

2 48

42

83

83

14

14

20

2.2

10.0

6 54

43

43

9

7

19

1.6

5.97

76

46

13

3

1

16

4.2

67

6.67

56

26

44

.4

10

2 1

92.1

65

4.

59

18

23

43.8

7

0 0

0

10.1

9 4.

61

10

.19

4.63

17.4

2 7.

95

-590

90

93

55

98

53

6.2

0.19

8 -1

085

72

89

53

83

32

1.1

0.00

3 9

-15

73

65

89

18

61

111.

2 0.

001

-2

0 60

41

90

1

35

8.6

0 0

88

88

0.53

0-5

76

76

78

54

24

0.32

1-1

071

61

74

46

23

16

0.

101

-15

69

51

74

15

8.8

10

0.01

2

-20

58

24

71

0 17

3.

6

98

98

91

101.

07

7336

.383

.2

0.39

6-5

94

94

91

45

94

100.

2 15

0.20

5 -1

083

83

91

47

98

81

.2

0.11

2-1

580

76

90

25

86

0.

001

-2

0 71

59

91

0

62

9.0

0 0

96

96

93

175.

67

7417

.432

4.5

0.52

1-5

92

92

99

87

87

0.31

4 -1

085

76

95

78

78

12

0.

064

-15

81

70

98

12

9.2

28

0.00

6

-20

72

65

96

0 0

6.26

0

0

94

94

3.4

19

250.

8 0.

399

-589

89

76

28

3.

6 19

19

5.0

0.09

8 -1

083

78

64

9

9.1

30

137.

3 0.

001

-15

78

94

9 4.

2 22

0.

001

-2

0 69

43

1

53

3.6

C

D fo

r A

4.50

43.4

15.

104.

494.

56

3.01

6.09

4.49

10.1

95.

514.

610.

084.

61

CD

for

B

4.50

43.4

15.

104.

494.

57

3.01

6.12

4.49

10.1

95.

514.

630.

084.

63

CD

for

A x

B

7.65

97

.03

11.0

7.

65

7.86

5.

14

10.4

37.

65

17.4

2 9.

45

7.95

0.

015

7.95

KHERA AND SINGH 215


provenances were considerably tolerant as is evident from their germination behaviour. No significant decline was observed in germination even up to -20 bars in P1, P3 and P4. However, in P5 the threshold was between -5 and 10 bars, and in P2, germination reduced significantly even at -5 bars.

In A. lebbek, germination percent and GV increased at a stress of -5 bars in all provenances except P4. The increase was maintained up to -10 bars in P5 and P3. P4 was the least tolerant provenance as, germination declined sharply even at a slight stress level of -5 bars. Seeds of D. sissoo and T. grandis were extremely sensitive to even a slight increase in stress level, except P3 where a stress of -5 bars enhanced percent germination and GV of D. sissoo seeds significantly.

Commencement of germination was delayed at higher levels of water stress in all the species except P3, P4 and P5 provenances of A. lebbek, and P4 and P5 provenaces of D. sissoo.

Wider response breadth of A. catechu, A. nilotica and A. lebbek shows their tolerance to increased stress levels. P1 and P3 provenances of D. sissoo also exhibited wider response breadth for stress tolerance (Table 3).

Across provenances, seeds of A. catechu, D. sissoo and T. grandis were highly sensitive to even a slight increase in the level of water stress as both germination percentage and germination value declined as the stress increased (Fig. 4). In A. nilotica and A. lebbek, however, germination percentage increased at moderately high levels of stress and afterwards in declined. This decline was significant only after –15 bars.

The decline in percentage germination in A. catechu, D. sissoo and T. grandis with increased levels of stress in the present study was consistent with the findings of Bokhari et al. (1975), Falleri (1994), Kaufmann & Eckard (1977) and Barnett (1969).

Reduction in seed germination at higher levels of water stress may be attributed to the moisture deficit in the seeds below the threshold which may lead to degradation and inactivation of essential hydrolytical and other groups of enzymes as suggested by Wilson (1971).

Among the species, A. nilotica and A. lebbek were found to be the tolerant to quite high levels of water stress. Bewley & Black (1985) also found in their experiments that the range of response to water stress was wide among species, from the very sensitive to the resistant. Resistant seeds may have an ecological advantage in that they can establish plants in areas in which drought-sensitive seeds cannot do so. Germination of these

two species, in the present study, was even benefited by slightly high levels of water stress.

It was observed that species with small sized seeds, viz. A. catechu and D. sissoo exhibited higher sensitivity and least tolerance to increased water stress than species with larger seeds, viz. A. nilotica and A. lebbek. This was in conformity with Rao & Singh (1989) who found that the inhibitory effect of water stress was markedly higher on the germination of smaller seeds than on larger seeds. However, in the present study T. grandis also exhibited lesser tolerance to the inhibitory effect of the water stress despite its bigger size.

Seeds of D. sissoo, A. nilotica and Albizia lebbek from different provenances responded diff-errntly to the inhibitory effect of water stress. Falusi et al. (1983) have also stated that intraspecific variations of drought tolerance can be quite marked, at least in species with a long distri-bution. Saint–Clair (1986) and Calamassi et al. (1980) have also demonstrated the existence of an early geographic variability between provenances and selection of those best adapted to water deficits.

Conclusions The differential germination response of

different provenances of the species to different environmental conditions suggests that a careful selection of provenances, for germplasm collection, is required in afforestation programmes. Variation among provenances was conspicuous in A. lebbek. Among the four environmental factors studied, intraspecific variation was more pronounced for water stress. Different provenances of all the five species varied in their response to increased level of stress. While some provenances were very sensitive to even a slight increase in the level of water stress, other emerged as almost resistant to high levels of water stress. Seeds from these provenances may be used for restoration of dry and degraded lands.

Acknowledgements

Authors are thankful to Dr. B.P. Singh, Head of the Department, for providing necessary facilities, and Dr. A. K. Saxena for his valuable suggestions. The first author is thankful to the Indian Council of Forestry Research & Education, Dehradun, for financial support.

KHERA AND SINGH 217

References Aldouse, J.R. 1972. Nursery Practice. Forestry Common

Bulletin. No. 43, London. Barnett, J.P. 1996. Moisture stress effects on

germination in longleaf and slash pine seeds. Forest Science 15: 275-276.

Bernstein, L. 1975. Effects of salinity and sodicity on plant growth. Annual Review of Phytopathology 13: 295-312.

Bewley, J.D. & M. Black. 1985. Seeds: Physiology of Development and Germination. Plenum Press, New York.

Bokhari, U.G., J. S. Singh & F.M. Smith. 1975. Influence of temperature regimes and water stress on the germination of three range grasses and its possible ecological significance to a shortgrass praire. Journal of Applied Ecology 12: 153-163.

Calamassi, R., M. Falusi & A. Tocci. 1980. Variazione geografica e resistenza a stress idrici in semi di Pinus halepensis Mill., Pinus brutia Ten. E, Pinus elderica M,edw (Geographical variation and resistance to hydric stress in seeds of Pinus halepensis, P. brutia and P. elderica). Annali dell’ Instituto Sperimentale per la selvicoltura 11: 193-230.

Catalan, L. 1992. Laboratory germination conditions for seeds of Prosopis flexuosa D.C. and P. chilensis (Molina) Stuntz. Seeds Science & Technology 20: 289-292.

Czabator, F.J. 1962. Germination value: an index combining speed and completeness of pine seed germination. Forest Science 8: 386-396.

Erasmus, D.J. & J. Van Staden. 1986. Germination of Chromolaena odorata (L.) K & R. achene: effect of temperature, imbibition and light. Weed Research 26: 75-81.

Falleri, E. 1994. Effect of water stress on germi-nation in six provenances of Pinus pinaster Ait. Seed Science & Technology 22: 591-599.

Falusi, M., R. Calamassi & A. Tocci. 1983. Sensitivity of seed germination and seedling root growth to moisture stress in four prove-nances of Pinus lahepansis Mill. Silvae Genetica 32: 4-9.

Ford-Robertson, F.C. 1971. Terminology of Forest Science and Products. Multilingual Forestry Terminology Series No 1 Society of American Forestry, Washington, D.C.

Gulliver, R.I. & W. Heydecker. 1973. Establishment of

seedlings in a changeable environment. pp. 433-462 In: W Heydecker (ed.) Seed Ecology. Butterworths, London.

Helmerick, R.H. & R.P. Pfeifer. 1954. Differential varietal response of winter wheat germination and early growth to controlled limited moisture conditions. Agronomy Journal 46: 560-562.

ISTA. 1985. International rules for seed testing rules and annexures. Seed Science & Technology 13: 299-519.

Kaufmann, M.R. & A.N. Eckard. 1977. Water potential and temperature effects on germination of Engelmann spruce and lodgepole pine seeds. Forest Science 15: 275-276.

Kozlowski, T.T. 1970. Physiological implications in afforestation. Proceedings of Sixth World Forestry Congress, Madrid 2: 1304-1316.

Levins, R. 1968. Evolutions in Changing Environment. Princeton, Princeton University Press.

Osmond, C.B., O. Bjorkman & D.S. Anderson. 1980. Physiological Processes in Plant Ecology towards a Synthesis with Atriplex. Sringler-Verlag Berlin.

Purohit, M. Jamaluddin & J.P. Mishra. 1998. Studies on germination and seed-borne fungi of some forest tree species and their control. Indian Forester 124: 315-320.

Rana, R.S. 1977. Plant adaptation to soil salinity and alkalinity. Proceedings of Indo-Hungarian Management of Salt Affected Soils (SSRI), Karnal 177-191.

Rao, P.B. 1986. Seed germination of Quercus leucotrichophora A. Camus ex. Bahadur and Pinus roxburghii Sarg on certain single factor environmental gradients. Proceeding of Indian Academy of Science (Plant sciences) 96: 63-69.

Rao, P.B. & S.P. Singh. 1989. Germination of certain climax and successional Himalayan trees as affected by moisture gradient: Implication for revegetation of bare areas. Tropical Ecology 30: 274-284.

Saint-Clair, P.M. 1986. Germination of Sorghum bicolor under polyethylene glycol induced stress. Canadian Journal of Plant Science 56: 21-24.

Uhvits, R. 1946. Effects of osmotic pressure on water absorption and germination of alfalfa seed. American Journal of Botany 33: 278-285.

Wilson, A.M. 1971. Amylase synthesis and stability in crested wheatgrass seeds at low water potentials. Plant Physiology 48: 541-546.

Germination of some multipurpose tree species in five ... · insolação, da temperatura, substrato...

Documents

Transcript of Germination of some multipurpose tree species in five ... · insolação, da temperatura, substrato...