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EFFECT OF TEMPERATURE ON CROP PRODUCTION

EFFECT OF TEMPERATURE ON CROP PRODUCTION A. Distribution of Plants : The most influential factors in the climate are temperature and moisture. Plants can grow only within certain limit of temperature. For each species and variety there are not only optimal temperature limits, but also optimal temperatures for different growth stages and functions, as well as lower and upper lethal limits. Temperature determines which species can survive in a particular region. Fig 1. Effect of temperature on the metabolic rate. Based on the temperature requirement vegetation can be classified as followsTable 2. Classification of vegetation based on temperature
Class Region Temperature Type of vegetation Megatherms Equatorial and tropical High temperature throughout the year Tropical rain forests Common crops cultivated Tropical crops like rubber, Cassava, Varieties of rice, etc. Megatherms Tropical and subtropical High temperature alternating with low temperature of winter Microtherms Temperate and high altitude plants (upto 12,000 ft of tropical and subtropical) Microtherms Aretic and Alpine regions (above 16,000 ft in tropics and 12,000 ft in temperature regions) Very low temperature Alpine vegetation Pines, spruce, etc. Low temperature Mixed coniferous forests Temperate crops like wheat, oats, potato etc. Tropical deciduous forests Sub-tropical crops like maize, sorghum, etc.

2. Germination : Temperature greatly influences germination of seeds. The inbibition phase of germination is primarily physical and shows low sensitivity to temperature. The subsequent phases of

germination (hydration and enzyme activation, enzymatic degradation of storage material, synthesis and growth of embryo) are temperature dependent due to biochemical processes involved. Cardinal temperature of different crops seed germination : Crops Seed Minimum Rice Maize Wheat Soybean Barley 11 9 4 9 4 Temperature ( OC ) Optimum 32 33 25 30 22 Maximum 41 42 32 41 36

Note : Cardinal Temperature : Cardinal temperature are those at which physiological process continue at such a rate so that it becomes critical for survival, growth and reproduction of plant. Or For survival each plant has a minimum, optimum and maximum temperature which is called cardinal temperature. 3. Photosynthesis : The effect of temperature on the rate of photosynthesis is little than on other processes. Very high and very low temperatures effect the photosynthetic rate adversely. According to mathaei (1904) the rate of photosynthesis increases with the rise in temperature from 50 to 370C beyond which there is a rapid fall. Between 50 to 370C, the rate of photosynthesis is doubled for every 100C increase in temperature. In desert and C4 plants photosynthesis may run relatively at high temperature. 4. Respiration : The rate of respiration increases with the rise of temperature up to a certain level, but beyond the optimum limit the respiration rate shows marked decrease. The rate of respiration becomes doubled at the increase of 100C above the optimum temperature provided other factors are favorable. In fact, respiration continues to increase after gross photosynthesis has reached its limitation by CO2 diffusion and enzyme kinetics. Thus the net photosynthesis is reduced (Fig. 3 ) 5. Flower Initiation :

In many crops (such as sugar beet, wheat) low temperature is required for flower initiation. The phenomenon of acquisition or acceleration of the ability to flower by chilling / low temperature treatment is termed as vernalization. 6. Induction of sterility : The occurrence of various cytological abnormalities in meiosis during the formation of the generative cell has been thought to constitute the primary cause of spikelets sterility. These abnormalities due to cold injury at the boot stage of rice plant. Sterility can also occur due to loss of stickiness of stigma and consequently failure of fertilization. 7. Growth Duration : Crop growth is influenced by temperature as every plant has its specific degree-day. A degree-day or heat unit is the departure from the mean daily temperature above the minimum threshold temperature. The minimum threshold is the temperature below which no growth take place. The threshold varies with different species of plants and for majority ranges from 4.50 to 12.50C. The growing degree-days are calculated with an equation written as : Growing degreedays (GDD) = Where, T max = Maximum daily temperature T min = Minimum daily temperature T t = Minimum threshold temperature It is important for the forecast of crop harvest dates, yield and quality. 8. Pest and Diseases : Usefully high temperature promotes the growth of weeds, insects and pathogens. 9. Temperature injury to plants : a. Low temperature injury : i. Chilling : Plants of tropical or subtropical, if exposed to low temperature above freezing point for some time, are found to killed or injured severely. ii. Heaving : Injury to plants is caused by a lifting upword of the plant along with the soil from its normal position. iii. Glaze : The weight of the ice coating on tree limbs cause them broken. iv. Suffocation : During winter the ice or the snow form a thick cover over the ground and the crop suffers from want of 02 v. Freezing : Water is frozen in the intercellular space of cell. b. High temperature injury :

i. Starvation injury: Shortage of food reserve for respiration. ii. Biochemical lesion : Break down of essential compounds like vitamins. iii. Toxin injury: Anaerobic respiration produces different compounds causing toxicity to plants. ATMOSPHERE: The atmosphere is a deep blanket of gases and suspended liquids and solids that entirely envelops the earth. Structure of Atmosphere : There are five concentric layers within the atmosphere which can be distinguished on the basis of temperature. These are as follows1. Troposphere 2. Stratosphere 3. Mesosphere 4. Mesosphere 5. Thermosphere, and 6. Exosphere 1. Troposphere : The lowest layer of atmosphere in which man and other living organisms live is called troposphere. It is about 20 KM in the equator and to KM in the polar region. It is a mixture of gases whose proportion is fairly constant. The composition of troposphere excepting water vapor and dust particles is as followsConstituent Percentage by volume a. b. c. d. e. f. g. h. i. j. k. l. Nitrogen Oxygen Carbon dioxide Argon Neon Helium Methane Nitrogen Oxide Krypton Hydrogen Xenon Ozone 78.0841 20.9476 0.0318 0.2340 0.001818 0.000524 0.0002 0.00005 0.000114 0.00005 0.0000087 0.000001

Troposphere is characterized by steady decrease in temperature and it may decrease up to 600C in the upper layer. It is the region of clouds, storms, and convective motion. The upper layer is called tropopause. [ The temperature of the air decrease by about 10C for every rise in 164.4 meters in altitude.] 2. Stratosphere : The second layer of air mass extending about 30 KM above tropopause is called stratopause. In this zone the temperature shows an increase from a minimum of about 600C to a maximum of about 50C. The increase in temperature is due to ozone formation under the influence of ultraviolet rays of solar radiation O2 20 O2+ 0 O3 Ozonosphere acts as a blanket that reduces the cooling rate of earth as well as prevents the lethal effect of ultraviolet rays. 3. Mesosphere : It is the third layer of atmosphere next to stratopauss. It is about 40 KM in height. It is characterized by low atomospheric pressure and temperature. The temperature begins to drop from stratopause goes on decreasing with the increase in height and reaches a minimum of about-95oC at a level some 80 to 90 KM above the earth surface. The upper layer of mesosphere is termed as mesopause. 4. Thermosphere : Next to mesosphere is thermosphere which extends about 500KM above the earth surface and is characterized by steady temperature increase with the height from meso-pause. In this region ultraviolet rays and losmic rays cause iovization of molecules like oxygen, and nitric oxide. In ionosphere, molecules of grass are so coidely spaced that high frequency cudible sound are not carried out by their atmosphere. 5. Exosphere : The rest region of atomosphere above the thermosphere is called exosphere or outer space which lalks atoms except thase of hydrogen and helium. This extends up to 3290 KM from the earth. Exosphere has a very high temperature due to solar radiation. The earths magnetic field becomes more important than gravity in distribution of atomic particles in the xosphere.

Extreme temperature effects on crops


There are two major forms of extreme temperature stress on crops - heat and cold. An increase in global temperatures may have either or both of these two acute effects: more frequent high temperature stress and less frequent cold temperature stress. Increase in temperature will lengthen the effective growing season in areas where agricultural potential is currently limited by cold temperature stress. Thus, increased temperature will cause a poleward shift of the thermal limits to agriculture. This poleward shift will be especially important for crops such as rice that have tropical centres of origin and adaptation but are also grown in temperate latitudes during warm seasons. Global warming impact will be greater in the northern than southern hemisphere because there is more high-latitude area cultivated in the northern hemisphere. Table 6.3. Effect of whole-plant warming on the rate of total dry matter and nitrogen accumulation, between days 10 and 20 after anthesis, in the grains of four cultivars of wheat
Cultivar Treatment AUS 22645 Kite Sonora WW15 C W C W C W C W Rate of increase (mg/grain/day) Total dry matter N content 1.9409 0.03002 2.0707(107) 0.48+003(160) 1.72+15 0.027004 2.2817(133) 0.043003(159) 1.6518 0.034009 2.0619(125) 0.051009(150) 1.8920 0.037005 2.3720(125) 0.053+005(143)

Plants were grown at 21/16C and some (W) were warmed, between 10 and 20 days after anthesis, to 33/25C and then returned to 21/16C where they stayed until maturity. Control plants (C) were grown at 21/16C throughout. Values given are the means .e; values in parentheses are percentage of cotton values. (From Bhullar and Jennar (1985); reproduced with permission.) Increased temperature would also affect the crop calendar in tropical regions. In the tropics, however, global warming, though predicted to be of only small magnitude, is likely to reduce the length of the effective growing season, particularly where more than one crop per year is grown. In semi-arid regions and other agro-ecological zones where there is wide diurnal temperature variation, relatively small changes in mean annual temperatures could markedly increase the frequency of highest temperature injury. For example, canopy temperature is 10 to 15C higher in dryland cotton (Gossypium hirsutum L.) than in irrigated cotton (Figure 6.2). Thus, global warming would reduce dry matter accumulation in dryland cotton because of increased respiration, and reduced photosynthesis and cellular energy.

In India, the growing season for wheat is limited by high temperatures at sowing and during maturation. As wheat is grown over a wide range of latitudes, it is frequently exposed to temperatures above the threshold for heat stress. For example, rainfed wheat depends on soil moisture remaining after the monsoon rains recede in September. High maximum and minimum temperatures in September (about 34/20C), which adversely affect seedling establishment, accelerate early vegetative development, reduce canopy cover, tillering, spike size and yield. Hence, sowing is typically delayed until after mid-October when seedbeds have cooled, though much of the residual soil moisture may be lost. High temperatures in the second half of February (25/10C), March (30/13C) and April (30/20C) reduce the numbers of viable florets and the grain-filling duration. High temperature stress particularly reduces yield of wheat sown in December/January which is necessitated in some regions because of the multiple cropping system. The situation is similar for sorghum (Sorghum bicolor (L.) Moench) and pearl millet (Pennisetum glaucum (L.) R.Br.) which are exposed to extreme high temperatures in Rajasthan, India. After sowing, air and soil temperatures often exceed 40C and midday soil surface temperatures above 50C are common (Figure 6.5). Acute effects of high temperature are most striking when heat stress occurs during anthesis. In rice, heat stress at anthesis prevents anther dehiscence and pollen shed, to reduce pollination and grain numbers (Mackill et al., 1982; Zheng and Mackill, 1982). Clearly, many crops in tropical areas are already subjected to heat stress. If temperatures increase further, crop failure in some traditional areas would become more commonplace.

Long-term effects of high temperatures on crops


More important than acute effects of extreme temperature stress are the chronic effects of continuously warmer temperatures on crop growth and development. Chronic effects of high temperature include effects on grain growth discussed above. Record crop yields clearly reflect the importance of season-long effects on crop yields: crops generally yield the most where temperatures are cool during growth of the harvested component. Crop growth simulations show that rice yields decrease 9% for each 1C increase in seasonal average temperature (Kropff et al., 1993). This chronic effect of high temperature differs significantly from the acute effect of short-term temperature events, because seasonal temperature effects are mostly a result of effects on crop development. For most grain crops, there is much greater genotypic variation in thermal requirements for vegetative than for reproductive development. As long-term temperatures increase, grain-filling periods decrease, and there appears to be little scope to manipulate this effect through existing genetic variation within species. Figure 6.5. Diurnal temperature data recorded in Fatehpur, Rajasthan, India. (Latitude 27C 37'N) in June 1989). Each measurement is the mean value from

three thermocouples placed at either 5 cm depth of soil ( ): 0.5 cm depth of soil ( ): or 150 cm above the soil surface ( ). (From Howarth (1991); reproduced with permission.)

References
Abrol, Y.P., Bagga, A.K., Chakravorty, N.V.K. and Wattal, P.N. 1991. Impact of rise in temperature on productivity of wheat in India. In: Impact of Global Climatic Change on Photosynthesis and Plant Productivity. Y.P. Abrol et al. (eds.). Oxford & IBH Publishers, New Delhi. pp. 787-798. Ahrens, M.J. and Ingram, D.L. 1988. Heat tolerance of citrus leaves. Hort Sci. 23: 747-748. Al-Khatib, K. and Paulsen, G.M. 1990. Photosynthesis and productivity during high temperature stress of wheat genotypes from major world regions. Crop Sci. 30: 1127-1132. Amthor. J.S. 1991. Respiration in a future higher CO2 world. Plant Cell Environ. 14: 13-20. Asana, R.D. and Williams, R.F. 1965. The effect of temperature stress on grain development in wheat. Aust. J. Agric. Res. 16: 1-13.

Bagga, A.K. and Rawson, H.M. 1977. Contrasting responses of morphologically similar wheat cultivars to temperatures appropriate to warm temperate climates with hot summers: a study in controlled environment. Aust. J. Plant Physiol. 4: 877-887. Behboudian, M.H. and Lai, R. 1994. Carbon dioxide in 'Virosa' tomato plants; responses to enrichment duration and to temperature. Hort. Sci. 29: 1456-1459. PHOTOSYNTHESIS AND HIGH TEMPERATURE STRESS Variability in leaf photosynthetic rates within or between species is often unrelated to differences in productivity. Similarly, high photosynthetic rates at high temperatures do not necessarily support high rates of crop dry matter accumulation. The temperature optimum for photosynthesis is broad, presumably because crop plants have adapted to a relatively wide range of thermal environments. A 1 to 2C increase in average temperature is not likely to have a substantial impact on leaf photosynthetic rates. Further, there is a possibility that photosynthesis of crop plants can adapt to a slow increase in global average temperatures. Thus, global warming is not likely to affect photosynthetic rates per unit leaf area gradually or on a closed canopy basis over the next century. While photosynthetic rates were found to be temperature-sensitive in other crops, wheat and rice appear to be different. In wheat, no measurable differences were found in photosynthetic rates per unit flag leaf area or on a whole-plant basis in the temperature range from 15 to 35C (Bagga and Rawson, 1977). In rice, there is little temperature effect on leaf carbon dioxide assimilation from 20 to 40C (Egeh et al., 1994). Recent research has shown significant variation among wheat cultivars with respect to reduction in photosynthesis at very high temperature. Photosynthesis of germplasm adapted to higher temperature environments was less sensitive to high temperature than was germplasm from cooler environments (Al-Khatib and Paulsen, 1990). When this germplasm was grown under moderate (22/17C) and high (32/27C) temperatures in the seedling stage or from anthesis to maturity, there was a highly significant correlation between photosynthesis rate and either seedling biomass (r=0.943***) or grain yield of mature plants (r=0.807**). Genotypes most tolerant to high temperatures had the most stable leaf photosynthetic rates across temperature regimes or they had the longest duration of leaf photosynthetic activity after anthesis and high grain weights. The above relationship was exemplified by 'Ventnor' from the high temperature area of Australia and 'Lancero' from the high altitude area of Chile (Table 6.1). See Al-Khatib and Paulsen (1990). Despite observed negative effects of high temperature on leaf photosynthesis, the temperature optimum for net photosynthesis is likely to increase with elevated levels of atmospheric carbon dioxide. Several studies have concluded that CO 2-induced increases in crop yields are much more probable in warm than in cool environments (Idso, 1987; Gifford, 1989; Rawson, 1992, 1995). Thus, global warming may not greatly affect overall net photo

NIGHT RESPIRATION RESPONSE TO CO2 AND TEMPERATURE In tomato (Lycopersicon esculentum) (Behboudian and Lai, 1994) and cotton (Gossypium hirsutum) (Thomas et al., 1993), elevated CO2 increased dark respiration possibly because of increased carbohydrate accumulation in tissues. The latter has been shown to increase alternative pathway respiration as well (Amthor, 1991). Apparent dark respiration may decline under elevated CO2 if there is dark CO2 fixation or if elevated CO2 directly inhibits or inactivates respiratory enzymes as may occur through increased formation or carbamate (Wullschleger et al., 1994). Table 6.1. Mean weekly photosynthetic rate (fmol CO2/m2/s) and duration of photosynthetic activity (weeks, in parentheses), and grain biomass of two wheat genotypes grown at two temperature regimes
Seedlings after two weeks of treatment 22/17 C 32/27 C 8.6 7.0 7.9 4.6 From anthesis to maturity 22/17 C 32/27C 6.4(7) 5.3 (7) 4.1 (10) 3.1(7) Grain biomass (g/tiller) 22/17 C 32/27 C 0.93 0.8 0.43 0.28

GENOTYPE Ventor Lancero

Figures in parentheses give duration of photosynthetic activity in weeks from anthesis to physiological maturity. (Modified from Tables 2 and 3 of Al-Khatib and Paulsen (1990); reproduced with permission.) Few studies have successfully partitioned the effects of elevated CO2 on growth and maintenance respiration. Both components appear to decline, probably because of decrease in leaf protein levels which results in reduced construction and maintenance costs (Wullschleger et al., 1994). Elevated CO2 reduced maintenance respiration of Medicago sativa and Dactylis glomerata at lower temperatures (15 to 20C), whereas elevated CO2 reduced growth respiration of M. sativaat 20 to 30C and D. glomerata at 15 to 25C (Ziska and Bunce, 1993).

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Topic 26.1
Stomatal Conductance and Yields of Irrigated Crops
In the last few decades, the breeding of agricultural crops for higher yields has been very successful. Breeders usually select for high-yielding genotypes empirically, without particular attention to specific plant traits that might be conducive to higher yields. However, comparing old, low-yielding lines of any crop with advanced, high-yielding lines shows clearly that many morphological, physiological, and biochemical traits have been altered by the intense selection pressures for higher yields. These changes indicate that selection for high-yielding genotypes has generated indirect selection pressures on the altered traits. If we exclude effects of genes that regulate the expression of two or more unrelated traits (pleiotropic genes), the study of high-yielding lines might reveal specific traits and genes associated with the higher yield. With this information, breeders could explicitly select for yield-enhancing traits to improve yields further. Changes in plant traits associated with yield increases can be studied conveniently in a historical series of a crop. Historical series include successive commercial releases within a breeding program, with each member of the series representing an incremental increase in yield. In studies aimed at characterizing relationships between specific traits and higher yields, all the members of the series are grown together under the same conditions, and the traits of interest, such as plant architecture or photosynthetic rates, are measured in parallel with yields. Recent studies of this type focusing on historical series of Pima cotton (Gossypium barbadense) and bread wheat (Triticum aestivum) have shown a remarkable positive correlation between yield increases and increases in stomatal conductance (Web Figure 26.1.A) (Lu et al. 1998).

Web Figure 26.1.A Stomatal conductance has increased in parallel with agronomic yields in irrigated Pima cotton (Gossypium barbadense) selected for higher yields at a high temperature. The figure shows the relationship between lint yield and stomatal conductance in a historical series of Pima cotton grown in

Arizona. The abbreviations P32 and PS-1 through PS-7 designate successive commercial releases between 1949 and 1996. (From Lu et al. 1998.)

Pima cotton is a high-quality fiber cotton grown in hot environments under intensive irrigation. The historical series used in the Pima studies included eight members, released between 1949 and 1996, and encompassing a nearly threefold increase in lint yield. The yield increase attained in each commercial release was accompanied by a corresponding increase in stomatal conductance. As Web Figure 26.1.A shows, stomatal conductance increased by about 30 mmol m2 s1 for each 100 kg ha1 increase in yield. Genetic crosses between low- and high-yielding Pima strains have shown that stomatal conductance in Pima cotton has a clear-cut genetic component (Web Figure 26.1.B) (Percy et al. 1996). The studies with stomata of Pima cotton are the first to show a clear-cut genetic regulation of stomatal conductance proper, of the stomatal response to temperature, and of proton pumping rates of isolated guard cells (Lu et al. 1998).

Web Figure 26.1.B Frequency distribution of stomatal conductance in parental populations of the high-yielding Pima cotton line (P73), the old Pima line (P32), and their F1 and F2 progeny. (A) Nonoverlapping distribution of stomatal

conductance in the two parental populations. (B) Distribution of stomatal conductance in an F1 population from a cross between the two parents. (C) Distribution of stomatal conductance in an F2 population derived from F1 parents. (After Percy et al. 1996.)

In the absence of explicit selection for higher stomatal conductance in the Pima breeding program, what could be the nature of the indirect selection pressures that caused the increases in conductance paralleling the increases in yield? Higher stomatal conductance increases CO2 diffusion into the leaf and favors higher photosynthetic rates (see textbook Chapter 9). Higher photosynthetic rates could in turn favor a higher biomass and higher crop yields. Advanced Pima lines show a higher photosynthetic capacity than older, low-yielding lines, but photosynthetic rates measured in the same leaves used to measure stomatal conductance were not positively correlated with yields (Radin et al. 1994). Thus, higher stomatal conductance appears to favor higher yields by a mechanism not directly related to photosynthesis. Evapotranspiration at the leaf surface lowers leaf temperature (see textbook Chapter 9), and higher stomatal conductance enhances this leaf cooling. Optimal daytime temperatures for growth, photosynthesis, and reproduction in Pima cotton are below 30C, while afternoon air temperature in the hot Pima-growing areas often exceeds 40C. Evaporative cooling of the leaves thus reduces the gap between optimal growth temperatures and air temperature, and provides heat resistance. Studies with the Pima historical series showed that, because of leaf cooling, leaf temperatures were several degrees lower than air temperature, and leaf and canopy temperature were lower in the advanced, high-yielding lines than in low-yielding lines. The same relationship among yields, stomatal conductance, and leaf temperature was found in a historical series of bread wheat grown in the warm Yaqui Valley of northwestern Mexico (Web Figure 26.1.C) (Lu et al. 1998). These studies indicate that selection for higher yields in irrigated crops grown at high temperatures imposes indirect selection pressures for high stomatal conductance that lowers leaf temperature and appears to reduce deleterious effects of heat stress on critical flowering and fruiting stages, thus resulting in higher crop yields.

Web Figure 26.1.C The relationship between grain yield and stomatal conductance in a historical series of semidwarf bread wheat grown in Ciudad

Obregn, Mexico. The abbreviations H1 through H8 designate successive commercial lines released by the International Maize and Wheat Improvement Center between 1962 and 1988. (From Lu et al. 1998.)

If selection for higher yields generates indirect selection pressures for higher stomatal conductance, can breeders explicitly select for higher conductance and thus obtain lines with higher yields? Studies with Pima cotton have shown that selection of high-conductance F2 progeny from a cross between high- and low-conductance parents produces highconductance F4 lines with higher lint yields than low-conductance lines have (Radin et al. 1994; Ulloa et al. 2000). These experiments indicate that high stomatal conductance could be used as a selection trait for high yields in irrigated crops grown at high temperature (Barbour et al. 2000). The observed increases in stomatal conductance in crops grown at high temperature also have valuable implications for models developed to predict global climate changes, which are expected to occur because of increases in atmospheric CO2 (see textbook Chapter 9). Under atmospheric CO2 concentrations that are twice as high as the present ones, evapotranspiration would decrease over the continents and air temperatures would increase significantly over the tropical land masses, amplifying the changes resulting from atmospheric radiative effects (Sellers et al. 1997). However, these models have yet to take into consideration the effect of higher temperatures on the stomatal control of evapotranspiration. Incorporation of the stomatal response to temperature in climate change models is likely to improve the accuracy of the predictions generated by the models. These studies illustrate both the complex interactions between physical and biological factors in the biosphere, and the advantage of a thorough understanding of the physiological properties of plants for efforts aimed at increasing agronomic yields and at improving models that predict global climate changes.

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