Agricultural and Forest Meteorology, 31 (1984) 251--260 Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands
251
GROWTH BEHAVIOUR OF FIELD GROWN SUGARCANE VARIETIES IN R E L A T I O N T O E N V I R O N M E N T A L P A R A M E T E R S A N D S O I L MOISTURE STRESS S. VENKATARAMANA, S. SHUNMUGASUNDARAM and K.M. NAIDU*
Sugarcane Breeding Institute, Coimbatore-641 007 (India) (Received December 8, 1983; accepted February 20, 1984) ABSTRACT Venkataramana, S., Shunmugasundaram, S. and Naidu, K.M., 1984. Growth behaviour of field grown sugarcane varieties in relation to environmental parameters and soil moisture stress. Agric. For. Meteorol., 31: 251--260. Growth analysis has been used to evaluate various sugarcane varieties for their drought resistance capacity under field conditions. Net assimilation rate (NAR) and relative growth rate (RGR) were high during early growth phases, but declined with the age of the crop. Leaf area ratio (LAR) and leaf area index (LAI) were enhanced with crop growth under normal irrigation (Nw) while drought (D) caused reduction in the leaf area production and LAI in varieties Co 349, Co 453 and Co 997. Environmental parameters such as air and soil temperatures were positively correlated with NAR while negatively related to leaf area production. Relative humidity (RH), rainfall during the crop season and number of rainy days had positive influences on leaf growth and LAI. Specific leaf area (SLA) or specific leaf weight (SLW) did not show a regular behaviour in either normally irrigated or moisture stressed conditions. INTRODUCTION G r o w t h b e h a v i o u r and d r y m a t t e r a c c u m u l a t i o n studies have long been in v o g u e to assess t h e p h y s i o l o g i c a l basis o f yields in c r o p plants (Evans, 1975). Certain breeding p r o g r a m m e s (Wallace et al., 1 9 7 6 ) have also utilized the g r o w t h analysis p r o c e d u r e s in evolving i m p r o v e d c r o p cultivars. Sugarcane, an a n n u a l c r o p o f t h e tropics, passes t h r o u g h f o u r distinct phases o f g r o w t h , n a m e l y , g e r m i n a t i o n , tillering, g r a n d g r o w t h and m a t u r i t y (Van Dillewijn, 1952). A l t h o u g h g r o w t h d e t e r m i n a n t s such as N A R , R G R a n d L A I , etc. have been a n a l y z e d in t h e m a j o r i t y o f t h e c r o p plants (Koller et al., 1 9 7 0 ; Wilhelm and Nelson, 1 9 7 8 ; F a k o r e d e and Mock, 1 9 8 0 ) , t h e r e have been fewer a t t e m p t s t o d e t e r m i n e t h e g r o w t h p o t e n t i a l , p a r t i c u l a r l y u n d e r d r o u g h t c o n d i t i o n s . S u g a r c a n e o f t e n e x p e r i e n c e s m o i s t u r e stress d u r i n g t h e f o r m a t i v e phase w h i c h generally coincides with t h e s u m m e r m o n t h s (Naidu, 1 9 7 6 ) a n d h e n c e r e d u c t i o n in g r o w t h associated with yield decline is a c o n s t a n t c o n s t r a i n t in sugarcane agriculture. Thus, to i d e n t i f y d r o u g h t - r e s i s t a n t varieties based u p o n their g r o w t h b e h a v i o u r , a screening p r o g r a m m e was initiated with a few p o p u l a r and w i d e l y a d o p t e d sugarcane * Author for correspondence. 0168-1923/84/$03.00
© 1984 Elsevier Science Publishers B.V.
252 varieties grown under normal water and drought conditions. The emphasis was on relating the environmental factors and the growth attributes to assess varietal variation in their dr ought resistance capacity. MATERIALS AND METHODS Sugarcane varieties Co 285, Co 312, Co 349, Co 419, Co 453, Co 617, Co 740, Co 997, Co 1148 and Co 6806 were field grown at Sugarcane Breeding Institute Farm, Coimbatore ( l l ° N ) , India, during the cropping season of 1978--79 (Table I). A strip-plot design with t w o t reat m ent s (Normal irrigation and Drought) was adopted; each replicated three times. The t r e a t m e n t s were in columns and the varieties were placed in rows. Two bud setts, th ir ty eight per row of 6.2 m (in addition to two setts on either end of the row which served as borders) were wet-planted during the first week o f January, 1978. A split dose of N was given as basal and at earthing up stage (90 kg acre -1 ). Manual practices as well as 2,4-D spray were a d o p t e d for weeding at least twice during the crop season. Plots allocated for drought t r e a t m e n t received no irrigation during the formative phase (days 60--150), while c o n t r o l plots received scheduled irrigation {once in 10 days). The crop stand was free f r om significant disease or insect damage. M e a s u r e m e n ts
Replicated samples were taken at m o n t h l y intervals com m enci ng from day 60 until the harvest of the crop. Each sample consisted of the above ground plant parts in 0.45 m 2 land area. The main shoot and tillers of each plant were separated into stem, leaves and sheaths and the leaf area of each segm e n t was calculated by multiplying length and width, incorporating appropriate correction factors. When the a m o u n t of a sample was in excess, a p o r t i o n o f representative sub-sample was used for dry weight d e t e r m i n a t i o n and was extrapolated for a whole sample. All samples were oven dried (80°C) until a cons t a nt weight was reached. The growth parameters were individually calculated using the standard formulae NAR
= (W2 -- W1) In L 2 -- In L 1 ) / [ ( t 2 - - t l ) (L2 --L~ )]
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Average leaf area o v e r t h e p e r i o d (t2 - - t l )
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=
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= L / L a n d area
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= ( L l n W2 -- In W 1 ) / ( W 2 - - W l )
SLW
= leaf (dry) wt./leaf area
SLA
= leaf area/leaf (dry) wt.
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254 (W 1 and W2 represent initial and final dry weight; L~ and L2 are the initial and final leaf area over the period (t2 -- tl )). All data were statistically analyzed and the mean values over the stages were worked out along with standard error determinations. A simple t-test was used to compare the two treatments of each variety. Correlations between the growth attributes and the weather parameters were worked out stage-wise to identify the effects of changing environmental variables on crop production. RESULTS AND DISCUSSION Sugarcane crop produces much dry matter by virtue of its unique C4 photosynthetic potential (Kortschak et al., 1965; Hatch et al., 1966), in addition to a high water use efficiency (Alexander, 1973). Net production in sugarcane is thus basically controlled at plant level, while environment and water availability m o d i f y the production potential in agricultural systems. In the present experiment, N A R , which is singularly responsible for growth, was high during the tillering and grand growth period, but declined sharply with the age of the crop. The mean values indicated that varieties Co 6806, Co 312, Co 740, Co 453, Co 285 and Co 419 possessed high N A R during the formative phase under Nw condition, while moisture stress caused marked reduction in Co 349, Co 419 and Co 453 (Table II). Although R G R was decreased with age coupled with drought effect, the mean values were, however, high in Co 312, Co 6806 and Co 1148 while reduction due to drought was evident in Co 349, Co 453, Co 1148 and Co 419. L A R were high in Co 1148, Co 453 and Co 740, even under moisture stress. S L W and S L A did not show a uniform pattern in either of the treatments. Reduction in L A I was observed in Co 349 and Co 419 under drought, while leaf weight in relation to plant dry weight was low in varieties Co 349, Co 453 and Co 997. Correlations between weather parameters and growth characteristics (Table III) have indicated a few salient points. Air temperature and soil temperature possessed a positive correlation with N A R (except in Co 453 and Co 6806) while relative humidity, rainfall and number of rainy days were negatively correlated. R GR was lowered with rise in RH% but increased with high air and soil temperature. L A R was positively related to temperature while RH% had negative association. S L W and S L A were n o t uniform during crop growth under both Nw and D conditions. However, S L W in Co 6806 possessed a highly significant positive correlation with the number of rainy days. Leaf area production was positively correlated with RH%, rainfall and number of rainy days, but negatively correlated with mean temperature, soil temperature, and evaporation rates. Rainfall induced more leaf area production under drought, particularly in Co 285, Co 740, Co 1148 and Co 6806, and temperatures possessed a highly significant positive correlation with respect to leaf dry weight/plant dry weight.
255 TABLE II Growth parameters of sugarcane varieties grown under normally irrigated (Nw) or drought stressed (D) conditions. The values are the means over ten stages of observation, and figures in parentheses represent the standard errors of mean difference. Variety
Co 285 Co 312 Co 349 Co 419 Co 453 Co 617 Co 740 Co 997 Co 1148 Co 6806
Tr
Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D
NAR a
RGR b
( g d m -2 day)
(g g-1 day)
0.049 0.048 0.072 0.161 0.049 0.041 0.052 0.047 0.055 0.044 0.046 0.051 0.056 0.053 0.035 0.046 0.046 0.049 0.074 0.063
0.017 0.015 0.020 0.018 0.017 0.014 0.015 0.014 0.017 0.014 0.016 0.017 0.015 0.014 0.015 0.017 0.016 0.015 0.018 0.020
(0.02) (0.06) (0.02) (0.02) (0.02) (0.02) (0.02) (0.02) (0.03) (0.02)
Variety
Tr
SLW e
SLA f
Co 285
Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D Nw D
10.73 10.89 (0.62) 15.96 (1.17) 14.92 12.80 (1.2) 13.58 11.82 (0.88) 11.97 12.82(0.96) 12.81 12.48 12.06 (0.78) 11.77 12.56 (1.20) 12.08 11.95 (0.94) 13.69 (1.17) 13.17 14.77 (0.64) 14.69
0.095 0.085 0.067 0.071 0.080 0.088 0.088 0.088 0.082 0.083 0.082 0.086 0.080 0.077 0.087 0.087 0.075 0.072 0.069 0.069
Co 312 Co 349 Co 419 Co 453 Co 617 Co 740 Co 997 Co 1148 Co 6806
LAI c
(0.009) (0.008) (0.008) (0.007) (0.007) (0.007) (0.008) (0.007) (0.008) (0.008)
LAR d
(dm 2 g~l) 4.47 2.98 3.75 2.66 5.62 1.42 3.98 2.58 4.66 2.86 5.36 3.98 5.36 4.26 6.48 4.54 4.82 3.62 4.56 3.22
(0.95) (0.97) (1.11) (0,87) (1.01) (1.45) (1.32) (1.61) (1.24) (1.14)
0.27 0.31 0.22 0.24 0.30 0.34 0.23 0.31 0.26 0.46 0.28 0.32 0.25 0.40 0.32 0.37 0.32 0.47 0.23 0.24
(0.06) (0.05) (0.05) (0.05) (0.17) (0.07) (0.13) (0.06) (0.22) (0.06)
Average leaf area Leaf dry weight/ over 30 days (dm 2) Total dry weight (0.009) (0.006) (0.007) (0.007) (0.006) (0.006) (0.011) (0.006) (0.009) (0.003)
203.33 147.69 173.88 134.39 264.40 69.95 188.80 146.73 221.37 143.87 259.81 201.94 248.83 208.13 305.67 232.32 217.13 171.75 200.00 157.01
(39.17) (41.27) (45.47) (40.70) (42.31) (63.02) (54.24) (70.57) (50.01) (46.54)
0.29 0.34 0.33 0.36 0.36 0.45 0.29 0.38 0.31 0.39 0.34 0.39 0.35 0.36 0.37 0.44 0.34 0.38 0.30 0.37
(0.05) (0.05) (0.05) (0.06) (0.06) (0.05) (0.08) (0.05) (0.07) (0.06)
a N A R Net Assimilation Rate. b R G R Relative Growth Rate. C L A I Leaf Area Index. d L A R Leaf Area Ratio. e S L W Specific Leaf Weight. f S L A Specific Leaf Area.
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259 Varieties selected for this study belong to divergent groups which include high yield, high percentage of sucrose in juice, early/late maturing, and drought susceptible/resistant types. Hence the growth pattern was not similar in all the varieties. Among the growth attributes, NAR, the primary determinant of growth, includes both photosynthesis and respiration and is influenced by incoming solar radiation. Photosynthetic potential would be higher only during early crop establishment while respiration dominates the later part and hence decline in N A R was imminent. SLA is usually more "plastic" (Kvet et al., 1971) especially with respect to the environmental variables. In the present study, drought influenced the number of individual leaves rather than the relative proportion of leaf dry weight to total dry weight and hence low L A I values were observed due to drought. LAI was singularly influenced by stand density which in turn was regulated by the crop stand in addition to the water availability. Since sugarcane grows well in warmer climates, radiation coupled with temperature influences the growth during the early phase. Thus, during the m a t u r i t y phase the growth parameters did not linearly respond to the changing environmental variables. Relative humidity and number of rainy days positively influenced the leaf area production. It is obvious from this experiment that the varieties Co 312, Co 740, Co 1148 and Co 6806 possessed better drought resistance capacity, while Co 349 and Co 419 were typically more sensitive. The usefulness of this study would be to select an ideotype for drought conditions (Asana, 1965; Donald, 1968) which might be structured by accounting these growth parameters. For instance, a variety similar to Co 6806, Co 740, Co 285 with respect to NAR; Co 312, Co 617 and Co 6806 with respect to RGR, Co 740, Co 1148 for high LAI, need to be evolved so that reduction due to drought might be minimized. In addition, maintenance of tissue turgidity and minimal water loss through transpiration should be progressively built up in the plant at the cellular level to adapt to drought environments, which could be of practical significance.
REFERENCES Alexander, A.G., 1973. Sugarcane Physiology. Elsevier, Amsterdam, pp. 752. Asana, R.D., 1965. Ideal and reality in crop plants. I. Wheat. J. Indian Agric. Res. Inst., 3 : 63--68. Donald, C.M., 1968. The breeding for crop ideotypes. Euphytica, 17 : 385--403. Evans, L.T., 1975. The physiological basis of crop yield. In: L.T. Evans (Editor), Crop Physiology. Some Case Histories. Cambridge University Press, Cambridge, pp. 327-355. Fakorede, M.A.B. and Mock, J.J., 1980. Growth analysis of maize variety hybrids obtained from two recurrent selection programmes for grain yield. New Phytol,., 85: 393--408. Hatch, M.D., Slack, C.R. and Johnson, H.S., 1966. Photosynthesis by sugarcane leaves. A new carboxylation reaction and the path of sugar formation. Biochem. J., 101: 103--111.
260 Koller, H.R., Nyquist, W.E. and Chorush, I.S., 1970. Growth analysis of the soybean community. Crop. Sci., 10: 407--412. Kortschak, H.P., Hartt, C.E. and Burr, G.O., 1965. Carbon dioxide fixation in sugarcane leaves. Plant Physiol., 40 : 209--213. Kvet, J., Ondok, J.P., Necas, J. and Jarvis, P.G., 1971. Methods of growth analysis. In: Sestak, J. Catsky and P.H. Jarvis (Editors), Plant Photosynthetic Production, Manual of Methods. W. Junk (publishers). The Hague, pp. 343--391. Naidu, K.M., 1976. Annual Report, Sugarcane Breeding Institute, India. Van Dillewijn, 1952. Botany of sugarcane. The Chromica Botanica Co., Waltham, MA, pp. 371. Wallace, D.H., Peet, M.M. and Ozbun, J.L., 1976. Studies of CO2 metabolism in Phaseolus vulgaris. L. and application in Breeding. In: R.H. Burris and C.C. Black (Editors), CO2 Metabolism and Plant Productivity, University Park Press, Baltimore, MD, pp. 43--58. Wilhelm, W.W. and Nelson, C.J., 1978. Growth analysis of tall fescue genotypes differing in yield and leaf photosynthesis. Crop Sci., 18: 951--954.