CC BY
59
Journal of Stress Physiology & Biochemistry, Vol. 14, No. 3, 2018, pp. 49-58 ISSN 1997-0838 Original Text Copyright © 2018 by Saravanan
ORIGINAL ARTICLE
OPEN rr\ ACCESS
3
Gas Exchange Characteristics and Water Use Efficiency in Eucalyptus Clones
S. Saravanan
1 Silviculture, Forest Management and Agroforestry division, Tropical Forest Research Institute (Indian Council of Forestry Research and Education), RFRC (PO), Mandla Road, Jabalpur - 482021. Madhya Pradesh, India
*E-Mail: [email protected]
Received September 3, 2018
The Institute of Forest Genetics and Tree Breeding, Coimbatore, India functioning under the Indian Council of Forestry Research and Education, Dehara Dun, has a long term systematic tree improvement programme in Eucalyptus spp. aimed to enhancing productivity and screening of clones for site specific. In the process, twenty four clones of Eucalyptus spp. were studied for the physiological parameters and water use efficiency from the established clonal trials. It also provides valuable information for establishing plantations at different geographic locations. Considerable variations were observed when the selected 24 clones of Eucalyptus spp. were subjected to physiological and WUE studies. The clones of C-188, C-186, C-14, C-10, C-123 and C-19 are falls in one cluster and the water use efficiency values are lower when compared to other clones and these clones are ranked first for high water use efficient clones for better productivity. Further, the clonal variation in physiological parameters and water use efficiency are discussed in detail in this article and clones suitable for large scale planting with higher productivity and WUE.
Key words: Physiological parameters, Water use efficiency, Eucalyptus clones, Productivity
Eucalypts are among the most widely cultivated forest trees in the world. The major Eucalyptus growing countries are China, India and Brazil and the growth rates that routinely exceed 35 m3 ha-1 year-1. Eucalyptus shows a broad productivity response depending on species, clones and soils factors (Onyekwelu et al., 2011). Eucalyptus sp. has some of the highest net primary productivity rates up to 49 m3ha-1year-1 (Hubbard et al., 2010). The high biomass accumulation potential makes Eucalyptus sp. a good prospect for timber, wood products and carbon sequestration projects. The pulp and paper industry is one of the key industrial sectors contributing to the Indian economy. There are 759 paper mills in India with an operating capacity of 12.7 million tonnes and consumption at 11 million tonnes with 9.3 kg per capita consumption of paper. The demand of wood from forest or commercial plantation for timber, fuel wood, pulp and paper production is increasing each year at an alarming rate. Therefore, there is an urgent need for improvement in production of forest resources to meet the needs. Large scale Eucalyptus plantations have been raised on forest and farm lands, community lands and road / rail / canal strips in India. However, most of these past plantations had very large genetic variation, low productivity ranging from 6 to 10 m3ha-1yr-1 and poor returns because inferior seed used for raising most of the target oriented plantations (Lal, 1993). Clonal selection and deployment in Eucalyptus is receiving attention as an intensive forest management tool for increased wood production. Many wood based industries in particular, pulp and paper industries are involved in plantation establishment program using clonal forestry approaches in the recent past. Over 5 million hectares of Eucalyptus plantations have been established throughout India. As a fast growing, remunerative and consistently demanded industrial wood, Eucalyptus has witnessed an unfettered support in India. Eucalyptus clonal planting has been said to have advantages includes quick provision of benefits associated with fast growth, short rotation for production of pulp wood (about 70 t ha-1), ready marketing and other reasons. In the present day scenario, research organizations and paper and pulp industries are
involving in developing new Eucalyptus clones for higher productivity with wide range of adaptability. Institute of Forest Genetics and Tree Breeding (IFGTB), Coimbatore also started the tree improvement program in Eucalyptus and shortlisted clones based on productivity. But there is a gap in knowledge on physiological and nutritional aspects especially the water and nutrient use efficiency of productive clones. Therefore the present study was undertaken to assess and rank the clones through gas exchange characteristics and water use efficiency of Eucalyptus clones along with the commercial clones available in the market at present and the seed origin seedlings for comparison purpose. The findings of the study will help in screening the clones for higher water use efficiency to specific site suitability and also adds value for the particular clone at the time of commercial release.
MATERIALS AND METHODS
To carry out the gas exchange characteristics and water use efficiency study, Eucalyptus clones are selected as the experimental material. This includes 24 clones and two seed origin seedlings. Among the 24 clones, 16 clones are shortlisted by IFGTB and these clones are numbered from C-7 to C-196. For comparison purpose, 8 clones (6 ITC clones and 2 TNPL clones) and two seed origin seedlings (each one from Tamil Nadu Forest Plantation Corporation and IFGTB) are selected and named as check clone 1 to 10. The physiological parameters and derived parameters like intrinsic and instantaneous WUE, etc. in different Eucalyptus clones from the established clonal trials, the Portable Photosynthetic System (PPS) model LiCOR -6400XT was used to measure the physiological parameters like net photosynthetic rate, stomatal conductance, transpiration rate, etc. from the Eucalyptus clones in the established clonal trials The collected wood samples are dried and powdered for water use efficiency study through the Isotope-ratio mass spectrometry (IRMS). The Gas exchange parameters measurements were taken between 9.30 am and 12.00 noon under cloud free conditions. Intrinsic water use efficiency (Pn/gs), instantaneous water use efficiency (Pn/E) and intrinsic carboxylation efficiency (Pn/Ci) were also determined using the primary data. Observations
from 25 ramets per clone in 3 replications were recorded for all the physiological parameters in the Eucalyptus clonal trials established in 4 locations to study the WUE. The data obtained on gas exchange characteristics viz., net photosynthetic synthesis, stomatal conductance, transpiration rate, instantaneous and intrinsic water use efficiency and the soil nutrients were used to perform correlation, regression and other statistical analysis using SPSS® 21.0 version and Microsoft® Excel 2007 (Panse and Sukhatme, 1985).
RESULTS
I. Physiological parameters
Observations on physiological parameters viz., net photosynthetic rate, transpiration rate, stomatal conductance, intercellular CO2 level, intrinsic carboxylation rate, intrinsic water use efficiency and instantaneous water use efficiency were recorded by using portable photosynthetic system (LiCOR). The data were analysed for statistical significance by using the software SPSS version 21. The results showed that, there is a significant difference among the clones with reference to the all the physiological parameters in different clones and the seed origin seedlings.
A. Net Photosynthetic rate (^ molm-2s-1)
The highest net photosynthetic rate was recorded in the clone C-188 (16.59) followed by C-186 (15.85) and the clone check clone 4 registered the lowest rate of net photosynthetic rate (6.21) followed by check clone 8 (7.29) compared to the mean 11.53. In the case of net photosynthetic rate, seed origin seedlings are registered lower rate when compared to the clones of Eucalyptus. The seed origin seedlings registered an average rate of 7.50 when compared to the clonal rate of 12-13.
B. Stomatal conductance (molm-2s-1)
In the case of stomatal conductance, the lowest stomatal conductance was recorded in the C-188 (0.0293) and check clone 7 registered the maximum of 0.0513 compared to the mean of 0.0386.
C. Transpiration (mmolm-2s-1)
From the established clonal trials, transpiration rate recorded the lowest value of 2.84 (C-188) to the highest
value of 9.25 in check clone 10 compared to the mean of 5.32.
II. Intercellular CO2 level and Intrinsic Carboxylation rate
Intercellular CO2 level and Intrinsic Carboxylation
rate data collected and analysed statistically and the results showed that, there is a significant difference among the clones.
A. Intercellular CO2 level (Ci-^l l-1)
In the case of intercellular CO2 level presence, higher the level of the CO2 which helps in tree species
for effective nutrient use efficiency and high photosynthetic activity which facilitate the growth over a period. Clone C-188 recorded the maximum intercellular CO2 levels (276.07) and followed by C-186 (273.01). The lowest level of intercellular CO2 was found in the
clone in check clone 10 (138.17). Clones of C-188, C-186, C-19, C-111 and C-111 form one cluster and register the higher intercellular CO2 levels of more than the mean of 222.28. The check clones 10, 9, 1, 2 3 and 8 are recorded the lower the rate of the intercellular CO2 and form a group. The remaining clone recorded intermediate values and form the clusters of intermediate.
B. Intrinsic Carboxylation rate (^mol m-2 s-1).
The intrinsic carboxylation rate was recorded higher in the C-188 (0.073) followed by C-10 (0.071). The lowest rate of intrinsic carboxylation rate was found in the check clone 4 (0.045) followed by check clone 5 (0.049), compared to the mean of 0.054. The clones of C-188, C-14, C-186 and C-123 forms a single cluster and registered the intrinsic carboxylation rate of more than 0.058. The check clones 4, 5, 1, 2 and 3 are recorded the carboxylation rate ranged between 0.034 and 0.045.
III. Water Use Efficiency
A. Intrinsic water use efficiency
The simplest form to define the Intrinsic water use efficiency (WUEintr), the ratio of photosynthesis to stomatal conductance to water use. Higher the ratio
between the photosynthesis and the stomatal conductance showed that better water use efficiency and the clone C-188 recorded the maximum value of 547.52 followed by C-123 (490.03). The lower the intrinsic water use efficiency value was found in check clone 4 (134.71) followed by and check clone 7 (135.48) with the mean of 301.05. The clones of C-188, C-123,
Table 1: Physiological parameters of Eucalyptus clones.
C-10, C-14 and C-186 are recorded greater intrinsic water use efficiency value and form a single group having water use efficiency on a par. The lowest values are recorded in the check clones 4, 7, 6, 5, 3 and 8 and these clones were recorded the minimum value of intrinsic water use efficiency.
Clone no Net photosynthetic rate (Pn- ц mol m-2s-1) Stomatal conductance (gn- molm-2s-1) Transpiration rate (E- mmolm-2s-1)
C 7 9.32 a-b 0.0330 a-b 4.43 a
C 9 10.16 b-c 0.0379 b-c-d 3.80 a
C 10 14.30 d-e 0.0292 a 3.73 a
C 14 15.32 d-e 0.0300 a 3.17 a
C 19 13.93d-e 0.0336 a-b 3.21 a
C 63 14.99 d-e 0.0342a-b 3.34 a
C 66 13.81 d-e 0.0416c-d e 3.80 a
C 100 12.89 c-d-e 0.0420c-d-e 4.37 a
C 111 14.86 d-e 0.0332 a-b 3.67a
C 115 12.74c-d 0.0416 c-d-e 4.05 a
C 123 15.85 d-e 0.0291 a 3.01 a
C 124 14.68 d-e 0.0388 b-c-d 3.55 a
C 186 15.97 d-e 0.0325 a-b 3.06 a
C 187 14.60 d-e 0.0340 a-b 3.50 a
C 188 16.71 e 0.0273a 2.84 a
C 196 15.01 d-e 0.0441 d-e 3.42 a
Check 1 7.44 a-b 0.0330 a-b 8.10 b-c
Check 2 7.23 a-b 0.0393 b-c-d 7.82 b-c
Check 3 7.53 a-b 0.0379 b-c-d 7.73 b-c
Check 4 6.00 a 0.0441 d-e 7.56 b-c
Check 5 7.14 a-b 0.0393 b-c-d 6.75 b
Check 6 8.44 a-b 0.0423 c-d-e 7.40 b-c
Check 7 7.07 a-b 0.0483e 6.90 b
Check 8 7.14 a-b 0.0292 a 7.90 b-c
Check 9 7.29 a-b 0.0300 a 8.80 b-c
Check 10 7.70 a-b 0.0336 a-b 9.12 c
Mean 11.47 0.0402 5.1
Table 2: Intercellular CO2 level and Intrinsic Carboxylation rates of Eucalyptus clones.
Clone no Intercellular CO2 (Ci-ц! l-1) Intrinsic Carboxylation (^mol m-2 s-1)
C 7 225.43 c-d-e 0.043a-b-c-d
C 9 252.91 e-f 0.047a-b-c-d
C 10 263.35 e-f 0.067d-e
C 14 232.44 d-e-f 0.069d-e
C 19 271.26 e-f 0.051a-b-c-d-e
C 63 264.93e-f 0.057c-d-e
C 66 248.53 e-f 0.057c-d-e
C 100 235.13d-e-f 0.057c-d-e
C 111 273.33 e-f 0.055b-c-d-e
C 115 242.229 d-e-f 0.054b-c-d-e
C 123 271.26 e-f 0.058c-d-e
C 124 245.43 d-e-f 0.051a-b-c-d-e
C 186 274.14e-f 0.058c-d-e
C 187 261.84e-f 0.056c-d-e
C 188 277.20 f 0.072e
C 196 271.25 e-f 0.055b-c-d-e
Check 1 179.85a-b 0.042a-b-c-d
Check 2 165.64 a-b 0.044a-b-c-d
Check 3 169.09 a-b 0.045a-b-c-d
Check 4 176.49 a-b 0.034a
Check 5 176.41 a-b 0.040 a-b-c
Check 6 185.43 a-b-c 0.046a-b-c-d
Check 7 199.46 b-c-d 0.036 a-b
Check 8 160.0 a-b 0.045a-b-c-d
Check 9 146.31 a 0.050a-b-c-d-e
Check 10 139.30 a 0.056c-d-e
Mean 223.41 0.052
Table 3: Water Use Efficiency in Eucalyptus clones.
Clone no Intrinsic Water Use Efficiency Instantaneous Water Use Efficiency WUE by Isotope Mass Spectrophotometer
C 7 282.42a-b-c-d 0.007 a-b-c 26.82 f-g
C 9 268.07a-b-c-d 0.010 b-c-d 26.73 f-g
C 10 489.73c-d 0.010 b-c-d 26.75 f-g
C 14 510.67c-d 0.011c-d 27.15 g
C 19 414.58c-d 0.012d 27.03 g
C 63 438.30a-b-c-d 0.010 b-c-d 26.57 e-f-g
C 66 331.97a-b-c-d 0.011 c-d 25.62 a-b-c-d-e
C 100 306.90a-b-c-d 0.010 b-c-d 25.01 a-b
C 111 447.59a-b-c-d 0.009 b-c 26.54d-e-f-g
C 115 306.25a-b-c-d 0.006a-b 24.79 a
C 123 544.67d 0.012d 26.78 f-g
C 124 378.35a-b-c-d 0.006a-b 25.38a-b-c
C 186 491.38b-c-d 0.011 c-d 27.12 g
C 187 429.41a-b-c-d 0.010 c-d 25.60 a-b-c-d-e
C 188 612.09d 0.013 d 27.21 g
C 196 340.36a-b-c-d 0.011 c-d 26.33 c-d-e-f-g
Check 1 225.45a-b-c 0.004 a 25.51 a-b-c
Check 2 183.97a-b 0.005 a 25.97 b-c-d-e-f
Check 3 198.68a 0.005 a 25.26 a-b
Check 4 136.05a 0.006 a 25.75 a-b-c-d-e
Check 5 181.68a-b 0.006 a-b 24.85 a
Check 6 199.53a-b 0.006 a-b 25.21a-b
Check 7 146.38a-b-c 0.007 a -b 25.62a-b-c-d-e
Check 8 244. a 0.004 a 25.28 a-b
Check 9 243.00a--b-c 0.003 a 25.56 a -b-c-d
Check 10 229.17 a-b-c 0.004 a 25.21 a-b
Mean 330.05 0.008 26.28
B. Instantaneous water use efficiency
Instantaneous water use efficiency is defined as the ratio of CO2 assimilation into the photosynthetic
biochemistry (A) to water lost, via transpiration through the stomata (T) or the ratio between the photosynthesis and transpiration. Increasing the external concentration of CO2 will increase instantaneous WUE, as the driving force for water loss will remain unchanged, while that for CO2 uptake will increase. Higher the ratio between the
photosynthesis and the transpiration showed that better the instantaneous water use efficiency. The clone C-188 recorded the maximum value of 0.013 followed by C-123 (0.012). The lower the instantaneous water use efficiency value was found in the check clones 9 (0.004) followed by check clone 10, 1 and 8 (0.004, 0.004 and 0.004) with the mean of 0.009. The clones of C-188, C-123, C-10, C-14 and C-186 are recorded the more the intrinsic water use efficiency value and form a single group in the case of high instantaneous water use
efficiency. The lowest values are recorded in the check clones of 9, 10, 1 and 8 and these clones were recorded the minimum value of intrinsic water use efficiency. C. Water Use Efficiency discrimination by Stable Isotope Mass Spectrophotometer
Carbon isotope discrimination has been used to assess the genetic variability in the driving force for CO2
uptake. Stable isotope discrimination delta (Д) has been used to assess genotype variation in WUE and physiological responses to environmental factors. Higher the value of the Д which indicates the better water use efficiency in the clones. The highest Д value of 27.21 was recorded in the C-188 clone followed by 27.15 in C-14, 27.12 in C-186 and 27.03 in C-19. The clone of C-115 registered the Д lowest value of 24.75 followed by 24.85 (in check clone 5) and 25.01 in C-100. The water use efficiency ranged between 24.75 and 27.15 among the clones. The grouping was done by using the DMRT and from the analysis, the clones of C-188, C-186, C-14, C-10, C-123 and C-19 are falls in one cluster and the water use efficiency for the Д values are higher when compared to other clones and these clones are ranked first for high water use efficiency. The check clones of 5, 4, 10, 9, 6 and C-100 are formed one cluster and registered the lower Д value and having lower water use efficiency.
DISCUSSION
Reports stated that photosynthetic rate varies among the plants belonging to different taxa and also among the varieties within the same species (Arora and Gupta, 1996). Olbrich et al. (1993) stated that, drought resistant clones may be selected for improving silvicultural practices for higher productivity and such clones need to be assessed physiologically under different conditions. Rekha Warrier et al. (2013) reported that, Clone EC 52 ranked top with reference to the net photosynthesis rate followed by EC 70. Clones EC 9, 10, 19 and 111 exhibited poor photosynthetic rates. The Pn values varied from 0.204 to 7.94pmol m-2 s-1 with a mean of 3.01pmol m-2 s-1 in different Eucalyptus clones. Net photosynthesis rate (Pn), is the important factor that determine the biomass production and Water Use Efficiency of a species. Variation in Pn, has been
reported as determinant of plant productivity in rubber (Nataraja and Jacob, 1999). Significant differences in Pn and stomatal conductance (gs) have been reported to exist in different tree species (Zipperlen and Press, 1996), viz., Eucalyptus camaldulensis (Farrel et al., 1996) Populus (Kalina and Ceulemans, 1997), Azadirachta indica (Kundu and Tigerstedt, 1998) and Hevea brasiliensis (Nataraja and Jacob, 1999). Considerable variation has been reported in clones of Eucalyptus camaldulensis for important physiological characteristics including high photosynthesis, carboxylation efficiency and water use efficiency (Warrier et al., 2010). Kannan and Venkatramannan (2010) studied the net photosynthetic rate in Eucalyptus clones and stated that, The Pn values varied from 10.05 to 37.80 pmol m-2 s-1 with a mean of 18.45 ± 6.70 pmol m-2 s-1. Kuyah Shem et al. (2009) reported that A, E and gs varied between species, being highest in Eucalyptus hybrid GC 15 (24.6 pmol m-2 s-1) compared to Eucalyptus hybrid GC 584 (21.0 pmol m-2 s-1), E. grandis (19.2 pmol m-2 s-1), C. africana (17.7 pmol m-2 s-1) and G. robusta (11.1 pmol m-2 s-1). The mean maximum net photosynthetic value for GC 15 (24.6 pmol m-2 s-1), GC 584 (21.0 pmol m-2 s-1) and E. grandis (19.2 pmol m-2 s-1) showed significant differences (P<0.001) in eucalyptus species. Balasubramanian et al. (2009) studied the photosynthetic rate in different Eucalyptus clones and revealed that, there is a significant difference among the clones under water logged conditions. Marrichi (2009) observed values from 25.7 to 31.6 pmol m-2 s-1 in eucalyptus plants at 16 months of age. White Head and Beadle (2004) found values of around 13-32 pmol m-2 s-1 for 11 species of Eucalyptus. Photosynthetic rate of any species is a direct indicator of plant growth and metabolism. Therefore, selection of a variety or species for a given geographical location could also be on the basis of its photosynthetic activities. A reduction in photosynthesis affected the accumulation of biomass in the plant.
The growth and development of trees on sites experiencing occasional periods of drought stress depends on the ability of stomata to control water loss while maintaining growth. Stomata also respond to CO2
as stomatal conductance decreases as CO2
concentration increases (Medlyn et al., 2001). The stomatal conductance decline in response to increase in CO2 concentration will, to some extent, compensate for the effect of increased CO2 on photosynthesis and may also reduce the transpiration rate and the integrated result of these effects is that an increase in atmospheric CO2 concentration generally increases water use efficiency (WUE) (Centritto et al., 1999). Stomatal conductance is of utmost importance when photosynthesis is concerned. Stomata play a pivotal role in controlling the balance between assimilation and transpiration. Rekha warrier et al. (2013) studied the clonal differences in stomatal conductance and the results showed that stomatal conductance varied between 0.008 to 0.264 mol-2 s-1 with a mean of 0.08 mol-2 s-1. The role of stomata in determining the water use efficiency is also well understood (Li, 2000). Kallarackal and Somen (1998) observed significant variations among different species of Eucalyptus viz. E. tereticornis, E. camaldulensis, E. urophylla, E.brassiana, E. pellita and E. deglupta in stomatal conductance. It was lowest in E. urophylla and highest in E. camaldulensis. As such, leaves must adjust their stomatal aperture to maximize photosynthesis while minimizing consequences of excessive water loss.
Intrinsic carboxylation efficiency was derived as the ratio of net photosynthetic rate to intercellular CO2
concentration (Pn/Ci). This result is in tune with Kannan and Venkatramannan (2010) who studied in Eucalyptus clones and stated that, among the 59 clones, EC 130
ranked first (197.30 pl l-1) for intercellular CO2 concentration. Clone EC 148 recorded the lowest value (92.58 pl l-1). The mean and standard deviation were 125.34 and 26.45 pl l-1 respectively. Among the eucalypt clones, EC 17-1, EC 1-7, EC 71, EC 72 and EC 130 recorded higher values for intrinsic carboxylation efficiency coupled with superior growth when compared to others. This ratio varied from 0.100 (EC 12-11) to 0.198 pmol m-2 s-1 (EC 17-1) with a coefficient of variation of 20 per cent (Kannan and Venkatramannan, 2010).
Water use efficiency is the ratio of CO2 assimilation
into the photosynthetic biochemistry (Pn) to water lost, via transpiration, through the stomata (E). Also defined as Water use efficiency is the ratio of A to transpiration (E), and is a measure of the amount of water used per carbon gain. A13C is related to the ratio of A/gs and termed intrinsic water use efficiency (Wi). Chunying Yin et al. (2005) found that there were significant interspecific differences in early growth, dry matter allocation, and WUE between two sympatric Populus species under well-watered and water-stressed treatments. Li (2000) reported that measurement of intrinsic WUE may be a useful trait for selecting genotypes with improved drought adaptation and biomass productivity under different environmental conditions. Higher intrinsic WUE was associated with productivity in Prosopis glandulosa and Acacia smallii (Polley et al.1996). It is reported that long-term structural and growth adjustments as well as changes in intrinsic WUE are important mechanisms of Acacia koa to withstand water limitation. Instantaneous WUE is estimated as the ratio of net photosynthetic rate to transpiration. Higher the value, better the efficiency of the plant to divert water for photosynthesis than transpiration. This result is in tune with Kannan et al. (2007) and revealed that, productive clones of Casuarina exhibited superior values of instantaneous WUE and the values ranged from 0.169 in CP 2401 to 0.477 pmol mmol-1 in CH-3004 clone. Similar findings have been reported for U. americana (Reich et al. 1989), Eucalyptus spp. (Sheriff, 1992), maritime pine (Guehl et al. 1995), Pinus radiata D. Don (Sheriff and Mattay, 1995), poplar clones (Liu and Dickmann, 1996), white spruce (Livingston et al. 1999) and Quercus robur L. (Welander and Ottosson, 2000). In these studies, the increase in intrinsic and instantaneous WUE was related to higher net photosynthetic rate coupled with low stomatal conductance and transpiration rate and resulted higher productivity.
Stable carbon isotope ratios (513C) in tree rings are the result of discrimination against the heavier 13CO2 during carboxylation and diffusion through the stomata, which are linearly related to the ratio of intercellular and
atmospheric CO2 (ci/ca). Therefore, time-integrated, intrinsic water-use efficiency can be inferred using stable carbon isotope ratios (513C) of plant tissues given its inverse linear relationship with Ci/Ca, whereby high water use efficiency is indicated by less negative 513C and low Ci/Ca and vice versa. Plant drought stress can be reflected to the degree by which plants discriminate against the heavier isotope carbon 513C during photosynthesis. Debbie Le Roux et al. (1996) reported that water use efficiencies differed significantly between clones and clonal variation in 513C is associated with variation in WUE. that water use efficiencies differed significantly between clones and clonal variation in 513C is associated with variation in WUE. Significant correlations between carbon isotope discrimination, instantaneous and growing season water use efficiencies were also found in western larch seedlings (Zhang and Marshall, 1993).
CONCLUSUSION
In the present study, the highest WUE value of 27.21 was recorded in the C-188 clone followed by 27.15 in C-14,-27.12 in C-186 and-27.03 in C-19. The clone of C-115 registered the lowest value of 24.75 followed by 24.85 (check clone 5) and 25.01 in C-100. The water use efficiency ranged between 24.75 and 27.15 among the clones. The grouping was done by using the DMRT and from the analysis, the clones of C-188, C-186, C-14, C-10, C-123 and C-19 are falls in one cluster and the water use efficiency values are lower when compared to other clones and these clones are ranked first for high water use efficient clones for better productivity. The above said clones exhibited superior growth coupled with favourable physiological characteristics including high photosynthesis, carboxylation efficiency and water use efficiency. Also, these clones are registered better intrinsic and instantaneous WUE and recorded the higher productivity. Further, these clones were tested for WUE by using the stable isotope mass spectrophotometer and the above said clones recorded higher values for better WUE positively correlated with higher productivity.
ACKNOWLEDGEMENT
The author is thankful to the Director General, Indian Council of Forestry Research and Education, Dehara Dun and the Director, Institute of Forest Genetics and Tree Breeding, Coimbatore, Tamil Nadu for providing the financial and other logistics supports to carry out the present study.
REFERENCES
Arora, D.K. and Gupta, S. (1996) Advances in Plant Phiosology, Vol 8. Anmol Publications Pvt. Ltd., New Delhi, 416 p.
Balasubramanian, A., Saravanan, K., Rajasekar, R. and Govinda Rao, M. (2009) Evolving superior genotypes of eucalyptus for biodrainage under water logged condition. 60th International Executive Council Meeting & 5th Asian Regional Conference, 6-11, December 2009, New Delhi, India.
Centritto, M., Magnani, F., Lee, H. S. J and Jarvis, P.G. (1999) Interactive effects of elevated CO2 and drougth on cherry (Prunus avium) seedlings: II. Photosynthetic capacity and water relations. New Phytologist, 141: 141-153.
Chunying Yin, Xiang Wang, Baoli Duana, Jianxun Luob and Chunyang Li. (2005) Early growth, dry matter allocation and water use efficiency of two sympatric Populus species as affected by water stress. Environmental and Experimental Botany, 53: 315322
Debbie Le Roux, William D. Stock, William J. Bond and David Maphanga. (1996) Dry mass allocation, water use efficiency and 513C in clones of Eucalyptus grandis, E. grandis x camaldulensis and E. grandis x nitens grown under two irrigation regimes. Tree Physiology, 16: 497—502. Farrel, R.C.C., Bell, D.T., Akilan, K. and Marshall, J.K. (1996). Morphological and Physiological comparisons of clonal lines of Eucalyptus camaldulensis: Response to drought and water logging. Aus.J. Plant Physio., 23: 497-507.
Guehl, J.M., Fort C. and Ferhi. A. (1995) Differential response of leaf conductance, carbon isotope discrimination and water use efficiency to nitrogen
deficiency in maritime pine and pedunculate oak plants. New Phytol., 131: 149-157.
Hubbard, R.M., Bond, B.J and Ryan, M.G. (2010) Evidence that hydraulic conductance limits photosynthesis in old Pinus ponderosa trees. Tree Physiology, 19: 165-172.
Kalina, J. and Ceulemans, R (1997) Clonal differences in the response of dark and light reactions of photosynthesis to elevated atmospheric CO2 in poplar. Photosynthetica, 33: 51-61.
Kallarackal, J. and Somen, C.K. (1998) Water relations and rooting depth of selected eucalypt species. KFRI Research Report 136.
Kannan C.S. Warrier, Ganesan, M and Venkataramnan, K.S. (2007) Gas exchange characteristics in Casuarina clones. Indian J. Plant Physiology, 12 (1): 83-87.
Kundu, S.K. and Tigerstedt, P.M.A. (1999) Variation in net photosynthesis, stomatal characteristics, leaf area and whole-plant phytomass production among ten provenances of neem (Azadirachta indica). Tree Physiol., 19: 47-52.
Kuyah Shem, Muthuri Catherine and Chin Ong. (2009) Gas exchange responses of Eucalyptus, C. Africana and G. robusta to varying soil moisture content in semi-arid (Thika) Kenya. Agroforest Syst, 75: 239-249.
Lal, P. (1993) Performance of Eucalyptus clones in Punjab. Proc.; National Symposium on Exotics in Indian Forestry, held at Department of Forestry and Natural Resources, PAU. Ludhiana, from March, 15 - 18, 2005, p. 45.
Li, C.Y. (2000) Population differences in water-use efficiency of Eucalyptus microtheca seedlings under different watering regimes. Physiol. Plant., 108: 134-139.
Liu, Z. and D.I. Dickmann. (1996) Effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones. Can. J. Bot., 97: 507-512.
Marrichi, A.H.C. (2009) Characterization of photosynthetic capacity and stomatal conductance in seven commercial clones of Eucalyptus and their
patterns of response to vapor pressure deficit. Dissertation (Master in Forest Resources) -School of Agriculture "Luiz de Queiroz", University of Sao Paulo, Piracicaba.
Medlyn, B.E., Barton, C.V.M., Broadmeadow, M.S.J., Ceulemans, R., De Angelis, P., Forstreuter, M., Freeman, M., Jackson, S.B., Kellomaki, S., Laitat, E., Rey, A., Roberntz, P., Sigurdsson, B.D., Strassemeyer, J., Wang, K., Curtis, P.S. and Jarvis, P.G. (2001) Stomatal conductance of forest species after long-term exposure to elevated CO2 concentration: a synthesis. New Phytologist, 149: 247-264.
Nataraja K.N. and Jacob, J. (1999) Clonal differences in photosynthesis in Hevea brassiliensis Mull. Arg. Photosynthetica, 36: 89-98.
Olbrich, B. W., Le Roux, D., Poulter, A. G., Bond, W. J. and Stock, W. D. (1993) Variation in water use efficiency and ö13C levels in Eucalyptus grandis clones. Journal of Hydrology, 150 (2-4): 615-633.
Onyekwelu, J. C., Stimm, B. and Evans, J. (2011) Review Plantation Forestry. In Günter et al. (Ed.), Tropical Forestry 8: Silviculture in the Tropics (pp. 399-454). Berlin: Springer-Verlag.
Panse, V.G. and Sukhatme, P.V. (1985) Statistical methods for agricultural workers, 4th Edition, ICAR, New Delhi.
Polley, H.W., Johnson, H.B., Mayeux, H.S. and Tischler, C.R. (1996) Impacts of rising CO2 Ecosystem Dynamics in a changing Environment. General Technical report. No. INT-GTR-338, pp. 189-194. Intermountain Research station, USDA Forest Service.
Reich, P.B., M.B. Walters and T.J. Tabone. (1989) Response of Ulmus americana seedlings to varying nitrogen and water status. Water and nitrogen use efficiency in photosynthesis. Tree Physiol., 5: 173184.
Rekha R. Warrier, Jayaraj, R.S.C. and Balu, A. (2013) Variation in Gas Exchange Characteristics in Clones of Eucalyptus сamaldulensis Under Varying
Conditions of CO2. Journal of Stress Physiology and Biochemistry, 9 (3): 333-344.
Sheriff, D.W. (1992) Nitrogen nutrition, growth and gas exchange in Eucalyptus camaldulensis, and Eucalyptus globulus seedlings. Aust. J. Plant Physiol. 19: 637-652.
Sheriff, D.W. and J.P. Mattay. (1995) Simultaneous effects of foliar nitrogen, temperature, and humidity on gas exchange in Pinus radiata. Aust. J. Plant Physiol., 22: 615-626.
Warrier, K.C.S. and Venkataramanan, K.S. (2010) Gas exchange characteristics in Eucalyptus clones. Indian Journal of Plant physiology. 15(3): 226-233.
Welander, N.T. and B. Ottoson. (2000) The influence of low light, drought and fertilization on transpiration
and growth in young seedlings of Quercus robur L. For. Ecol. Manage. 127: 139-151.
Whitehead, D. and Beadle, C.L. (2004) Physiological regulation of productivity and water use in Eucalyptus: a review. For. Ecol. Manage, 193: 113140.
Zhang, J. and J.D. Marshall. (1993) Population differences in water-use efficiency of well-watered and water-stressed western larch seedlings. Can. J. For. Res. 24: 92--99.
Zipperlen, S.W. and Press, M.C. (1996) Photosynthesis in relation to growth and seedling ecology of two dipterocarps rain forest tree species. J. Ecol. 84: 863-876.
