Agronomy Journal

Volume 93 MarchApril 2001 Number 2

SYMPOSIUM PAPERS
System Analysis of Plant Traits to Increase Grain Yield on Limited Water Supplies
Thomas R. Sinclair* and Russell C. Muchow ABSTRACT
Crop growth in commercial situations usually requires maximizing grain yield on limited available water resources, which results in maximizing the ratio of yield to evapotranspiration. A system analysis was undertaken to identify those plant traits that might be altered to improve crop yield in a water-limited environment. A mechanistic crop model was used to simulate maize (Zea mays L.) yield over 20 yr at Columbia, MO, which had high interannual variability in precipitation. Because sorghum [Sorghum bicolor (L.) Moench] is known to be better adapted to drier environments, a number of individual plant traits were adjusted in the maize model to represent a sorghum crop. For the tested environments, it was found that decreasing leaf size and increasing seed growth rate both resulted in decreased yield and a decreased ratio of grain yield to evapotranspiration. On the other hand, increasing the depth of water extraction resulted in increased yields and an increased ratio of grain yield to evapotranspiration. Combining sorghum-like traits in the maize model also increased yield and the ratio of grain yield to evapotranspiration when averaged for all years. For all seasons where simulated yield was less than approximately 550 g m2, grain yields were greater for a crop with sorghum-like traits than for a crop with maize traits.

here is a long history of research on water use efficiency (Tanner and Sinclair, 1983), and coincidentally, a myriad of expressions for water use efficiency (Sinclair et al., 1984). Increased water use efficiency is of greatest interest to growers when yields are maximized for the available water supply in each growing season. High water use efficiency in itself is of little interest if it is not associated with high yields. For example, plants that use Crassulacean acid metabolism as a precursor to CO2 assimilation have low stomata conductances during the daylight, and consequently have very high water use efficiencies but very low growth rates.

T.R. Sinclair, USDA-ARS, Agron. Physiol. and Genet. Lab., Univ. of Florida, P.O. Box 110965, Gainesville, FL 32611-0965; and R.C. Muchow, CSIRO, Cunningham Lab., Division of Tropical Crops and Pastures, 306 Carmody Rd., St. Lucia, Brisbane, QLD 4067, Australia. Received 28 Jan. 2000. *Corresponding author (trsincl@gnv.ifas. ufl.edu). Published in Agron. J. 93:263270 (2001).

Economic benefits from increased water use efficiency under water-limited conditions are usually achieved only if yield is maximized for the available water. There are, of course, several approaches to investigating plant traits for the purpose of increasing yields under water-limited conditions. One experimental approach is to examine the changes in plant traits that may have occurred in cycles of selection leading to increased yields under water deficit conditions. The difficulty in this approach is that there are likely to be a number of interacting and confounding factors. For example, Bolanos et al. (1993) found that few plant traits were correlated with the increase in yield under water deficit conditions observed through eight cycles of selection in a tropical maize population. Of the many traits studied, including osmotic adjustment, only those factors reflecting a reduction in the days to flower, which apparently allowed an escape of drought stress in this environment, were associated with the yield increase. In the approach used here, a system analysis is presented that focuses on specific plant traits that have been hypothesized as approaches leading to higher yields under water-limited conditions. The influence of individual plant traits are examined in a system analysis using a mechanistic crop model to simulate crop growth under a range of environments. The outputs from these simulations offer quantitative predictions of anticipated yields in response to changing any individual trait of the plant. In addition, the simulation of crop growth over a number of years offers a perspective on the yearto-year variability that is not sampled in conventional experimental approaches. System analyses of crop yield under water-limited conditions are not new. Previous analyses, however, examined the potential response in crop growth to alterations in only a few traits. Jordan et al. (1983) found that deeper rooting would clearly increase crop yield
Abbreviations: AMAX, area of the largest leaf on the plant; DHI, rate of harvest index increase; FTSW, fraction of transpirable soil water; HI, harvest index; RUE, radiation use efficiency; TU, thermal units.

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while earlier maturity and osmotic adjustment had little or no benefit. Similarly, Jones and Zur (1984) showed that an increased soil volume occupied by roots was the most effective adaptive mechanism for increasing growth during simulations of a 10-d drying cycle. Neither of these studies considered the possibility that water deficits may develop that result in the premature senescence of the crop. A simulation study by Sinclair (1994) on the influence of rooting depth on maize yield did include, however, the possibility of crop termination due to severe soil drying. For three locations in the USA, these simulations confirmed the advantage of deeper rooting across a number of years. Other simulation analyses of plant traits for water deficit conditions include those of Muchow et al. (1991) and Hammer et al. (1996). In contrast to the few plant traits that have been subjected to systems analyses, a large number of traits have been speculated as potentially contributing to increased crop productivity under water-limited conditions. Ludlow and Muchow (1990) discussed 16 such traits, and they recommended that eight of the traits were suited for intermittent stress conditions in modern agriculture. Their top three recommendations in order of priority were to match plant phenology to water supply, osmotic adjustment, and rooting depth. Because their priority list is inconsistent with the results of previous simulation analyses and because they listed a number of plant traits that were not previously considered in a simulation analysis, it is useful to reconsider the possible contribution of various plant traits to increasing yield under water-limited conditions. Consequently, this analysis was undertaken to examine in a maize model the putative advantage of a number of plant traits in a water deficit environment. The approach was to modify individual traits that were appropriate for maize so that they reflected the behavior of those traits in sorghum. Sorghum is generally felt to be more suited to dry climate conditions than maize. The traits that were modified included the depth of soil water extraction, individual leaf size, the rate of leaf appearance, CO2 assimilation activity, early stomata closure, delayed stomata closure, and grain growth rate and duration. In all cases, the crop was assumed to be vulnerable to crop termination due to severe water deficits. One trait that was not considered for adjustment was the transpiration efficiency coefficient used to calculate the daily crop transpiration rate based on the daily crop mass accumulation (Muchow and Sinclair, 1991). The transpiration efficiency coefficient for crop gas exchange is well defined and is relatively stable over a range of conditions (Tanner and Sinclair, 1983). Certainly, if this coefficient is less in a particular cultivar than might be expected for that species, then transpiration efficiency and likely yield can be increased by genetically increasing this coefficient toward the expected value. In this analysis, it will be assumed that the transpiration efficiency coefficient of the crop is stable at 9 Pa, which is near the maximum value for C4 species. Previously, this value for the transpiration efficiency coefficient was assumed to be 9 Pa for both maize and sorghum (Muchow and Sinclair, 1991; Sinclair et al.,

1997). However, maintaining a constant transpiration efficiency for crop gas exchange does not preclude substantial changes in the overall ratio of crop yield to total water loss because of the many factors influencing transpirational water loss, soil evaporation, and the formation of grain yield. A number of years were simulated because the weather scenario influencing the development of water deficits during the season can influence crop response. However, this analysis is illustrative because it was done for only a single location, and altered environmental conditions could well influence the sensitivity of crop growth and yield to any particular trait. It is anticipated that this example will illustrate the quantitative merits of specific plant traits for increasing yields under water deficit conditions. MATERIALS AND METHODS Model Description
The model used to analyze the crop response to alterations in various plant traits was the relatively simple mechanistic model for maize published by Muchow and Sinclair (1991). This model was shown to be robust in that simulated yields responded to variations in water availability in a manner similar to the observed responses. More recently, the same model structure was used to describe sorghum growth and yield, and the main modifications were in the coefficients used to describe the development of the sorghum plant (Sinclair et al., 1997). Again, the results of the model in predicting sorghum growth proved to be robust across environments with differing water availability. For many of the plant traits examined, the value of a variable describing the behavior of a trait in maize was simply changed to a value representative of sorghum. Each trait was singly changed in the maize model, and the resulting yields were compared with the nominal maize yields. A final set of simulations was done in which all of the putative traits were included for increased yields under water deficit conditions. Below are the specific traits that were examined for improving crop yield, and consequently, the ratio of grain yield to evapotranspiration. Depth of Soil Water Extraction Experimental evidence from sorghum has shown that the water extraction depth of a drought tolerant cultivar was at least 40 cm deeper than a cultivar that lacked tolerance (Salih et al., 1999). This trait was found in previous simulation studies to be particularly important in making more water available to the crop if water is stored at depth in the soil. In these simulations, the potential store of soil water is calculated as the depth of soil water extraction multiplied by 0.13, which is a reasonable estimate of the volumetric fraction of plant available soil water for many soils (Ratliff et al., 1983). The original maize simulations of Muchow and Sinclair (1991) assumed a store of plant available soil water of 135 mm, or a water extraction depth of approximately 100 cm, based on experimental observations. For the analysis presented here, the nominal rooting depth was assumed to be 80 cm (104 mm water). While this may be slightly less than would be expected of maize, it is not much less than that observed for maize by Carberry et al. (1989). The use of a minimal value for the extraction depth of maize allows for an increased occurrence in the development of stressed conditions in the simulations

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to emphasize the potential for trait changes to increase yields. Two sets of simulations were undertaken with an increased depth of soil water extraction: 100 and 120 cm (130 and 156 mm water). The results of Birch et al. (1990) showed in a side-by-side comparison that the depth of water extraction by sorghum was greater than that of maize. Leaf Size Salih et al. (1999) found that the leaf area produced by a drought tolerant cultivar of sorghum was only about 45% of that produced by a cultivar that lacked tolerance. In these simulations, the leaf area was decreased by changing the single variable (AMAX) that defined the area of the largest leaf on the plant. All of the other leaves on the plant are scaled based on the value of AMAX. Changes in AMAX simulated a change in the leaf area development and the overall potential in the crop leaf area index. In the maize model of Muchow and Sinclair (1991), AMAX was set equal to 750 cm2. A possibility for increasing yield on limited water is to decrease the leaf area so that soil water is conserved to avoid stress conditions, especially late in the season. The influence of this trait on yield was tested by changing AMAX to 500 cm2, which is representative of the smaller leaves usually produced by sorghum (Muchow, 1988a). Rate of Leaf Appearance Similar to the above approach, the rate at which leaves appear will influence the rate at which the canopy leaf area develops. An important consequence of changing only the rate of leaf appearance is that slowing this rate lengthens the duration of the vegetative development phase. The rate of maize leaf appearance (Muchow and Carberry, 1989) was changed to a function describing the slower appearance rate of sorghum leaves (Muchow and Carberry, 1990). Carbon Dioxide Assimilation Because transpiration is intimately linked to CO2 assimilation by the constancy in the transpiration efficiency coefficient, any change in CO2 assimilation results in a proportional change in water loss. In the model, CO2 assimilation is defined by the radiation use efficiency (RUE), which for maize, was equal to 1.6 g MJ1 total solar radiation (Muchow and Sinclair, 1991). Decreasing RUE results in decreased crop growth, and consequently, a decreased transpiration rate so that the development of a possible soil water deficit is delayed or avoided. The importance of this trait was tested by changing the RUE to 1.25 g MJ1, which is more representative of sorghum (Muchow and Davis, 1988). These assumed values for maize and sorghum are fully consistent with values reported by Kiniry et al. (1989) based on a summary of published results up to that point. Early Stomata Closure Radiation use efficiency is down-regulated in the model as soil water deficits develop, which are an expression of stomata closure. This down regulation occurs as a function of the fraction of transpirable soil water (FTSW). The value of the FTSW ranges from 1.0 at field capacity to 0.00 when all of the water available for transpiration has been extracted (the value of the FTSW can be 0.00 as a result of continued soil evaporation even though the transpiration rate is zero). Transpiration as a function of the FTSW has been described in a number of species, and a relatively stable exponential function has been obtained across a range of conditions (Sinclair et al.,

1998). Down regulation does not usually occur until the FTSW has decreased to a value of about 0.30 to 0.35. For example, the response of down regulation in maize does not result in an RUE decrease to a value that is 90% of the well-watered crop until the FTSW has decreased to a value of 0.29 (Eq. [1] of Muchow and Sinclair, 1991). To conserve soil water, down regulation might be induced at an earlier stage in soil drying. While there appears to be only small variation among crop species in the FTSW at which the down regulation is triggered, Ray and Sinclair (1997) did identify a cultivar of maize in which down regulation was triggered at a significantly higher FTSW than the other cultivars that were tested. Even though maize and sorghum have an equivalent stomata response to soil drying, the possibility of inducing early stomata closure was simulated. Early stomata closure was induced so that RUE was decreased to 90% of the well-watered value when the FTSW decreased to only 0.44. Delayed Stomata Closure An alternative strategy to the conservative response, in which down regulation is induced earlier in soil drying, is to sustain plant growth by delaying stomata down regulation. In this case, the crop continues to extract water at a high rate until the soil becomes very dry, i.e., there is a very low FTSW, in anticipation that there will be rain or irrigation before severe water deficits develop. This strategy, therefore, avoids the initial negative consequences of soil drying by continuing plant growth. This approach is one likely expression of osmotic adjustment in that the decrease in stomata closure is delayed (Ludlow and Muchow, 1990). While no difference between maize and sorghum for delayed stomata closure has been reported, this response was simulated in the model so that RUE did not decrease to 90% of the well-watered value until the FTSW reached 0.11. Grain Growth Rate and Duration A long period of grain growth results in soil water loss that could subject the crop to severe water deficits. Therefore, yield might be enhanced in water-limited situations if the rate of grain growth is increased and the duration of grain growth is shortened. For example, the data of Ehdaie (1995) showed that there was a negative association between yield and the length of the period from anthesis to maturity among eight wheat (Triticum aestivum L.) cultivars grown under waterdeficit conditions. In the model, the daily rate of grain growth is calculated based on a constant linear increase in the harvest index (DHI). A low DHI value results in a low estimate of the rate of grain growth. The DHI for maize is approximately 0.0150 while it is approximately 0.0185 for sorghum (Muchow, 1988b). Coupled with the difference in the DHI is a variation in the duration of grain growth, which is expressed in thermal units (TU). For maize, the length of the grain duration period was defined as 1150 TU (base temperature 0C), and for sorghum, the duration was defined as 800 TU (base temperature 0C). Combined Traits As a final test of the yielding capability under water-limited conditions of crops with sorghum traits in contrast to maize traits, leaf size, rate of leaf appearance, CO2 assimilation, and grain growth rate and duration were combined to convert the maize crop to one with sorghum-like traits. In this case, the adjustment in the depth of soil water extraction for the sorghum-like trait was increased from 80 to 100 cm. A key feature of all of these tests was the inclusion of a

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threshold soil water content at which the crop was assumed to be terminated. Therefore, crop survival of severe water deficit conditions was not assured in these analyses, and large decreases in yield could result from the termination of crop growth. In these analyses, the soil water content at which crop growth was terminated was set at a FTSW of 0.02 or less when the HI of the crop was 0.2. The HI restriction was included to allow a minimum production of crop yield even under the most extreme water deficits. The fatal FTSW of 0.02 was slightly less than the value of 0.00 that we previously assumed for maize (Muchow and Sinclair, 1991). Nevertheless, with an assumed soil depth of 80 cm, this threshold resulted in early crop termination in about half of the years in the tested environment.

Simulated Environment
The results of a system analysis, obviously, depend on the environment for which the simulations are done. For this analysis, weather data for Columbia, MO were used because of the high interannual variability in precipitation of this location during the growing season. The mean 20-yr annual rainfall during the growing season for 1969 to 1988 was 411 mm, but the range in growing-season rainfall was 169 to 772 mm. It was assumed that the soil water profile was fully charged at the beginning of each growing season. At crop emergence, the depth of soil water extraction was taken to be 30 cm. Because it was assumed that the volumetric faction of plant available soil water was 0.13, the initial store of available soil water was 39 mm. The depth of soil water extraction was increased by 3.35 cm d1 (Robertson et al., 1993) until the maximum depth was achieved. The sowing date in each year was held at 23 April. Crop emergence was simulated to occur at 87 TU following sowing. The plant population was held constant throughout the simulations at 7 plants m2. Also, the number of leaves per plant was maintained at 18.3 for all simulations. For each year, the yield for a simulation with an altered trait was compared with the simulated yield for the nominal maize crop. The difference between the yields was calculated for each year and plotted against the nominal yield for that year. Such a presentation highlights the range of yield changes simulated when a trait is altered from the nominal case.

Fig. 1. Plot for each of 20 yr of simulated maize yield against precipitation during the growing season at Columbia, MO.

was highly variable from year to year (Fig. 1) such that a quantitative evaluation of traits is unreliable based solely on precipitation. There were 11 seasons where the FTSW reached levels that resulted in the termination of maize growth in the simulations. As discussed later, the sensitivity in these simulations resulted, in part, because the depth of soil water extraction was assumed to be only 80 cm. In 4 out of 20 yr, crop termination resulted in yield losses 300 g m2. Although the severity of stress in this environment and an assumed 80-cm depth of water extraction in the simulations did not favor maize production, these conditions offered a good opportunity for examining the putative benefits of changing various plant traits to increase crop yield.

Depth of Soil Water Extraction

The benefit of an increased rooting depth in avoiding early crop senescence is illustrated by comparing the simulated FTSW through the 1982 growing season for the 80- and 100-cm rooting depth (Fig. 2). In this year, there was a dramatic decrease in the FTSW that began at about 80 d after sowing. The crop had a high rate of transpiration during this period, and there was virtually no precipitation to recharge the soil water. In the simulation for maize with an 80-cm rooting depth, the FTSW continued to decrease to the fatal level on Day 103 with a predicted yield of 269 g m2. On the other hand, with a 100-cm depth of water extraction, the fatal FTSW was

RESULTS
The variation in weather among years at Columbia, MO resulted in simulated grain yields for maize ranging from 87 to 976 g m2 dry weight and a mean yield of only 409 g m2 (Table 1). The simulated yield response
Table 1. Summary of mean values for yield, transpiration, evapotranspiration (ET), and the ratio of yield to ET from 20 yr of simulations with changes in various crop traits.
Trait Maize 100-cm Rooting Depth 120-cm Rooting Depth AMAX 500 cm2 Slow leaf development RUE 1.25 g MJ1 DHI 0.0185 Early stomata closure Delayed stomata closure Sorghum-Like Yield g m 2 409 507 563 370 540 421 368 445 381 474 Transpiration mm 171 201 225 150 215 166 158 173 171 184 ET 387 419 442 389 408 386 366 391 383 402 (Yield/ET) mg g1 0.99 1.16 1.23 0.90 1.23 1.04 0.96 1.08 0.93 1.14

RUE, radiation use efficiency. DHI, rate of harvest index increase.

Fig. 2. Plot of the fraction of transpirable soil water (FTSW) against days after sowing for simulated maize crops in 1982 with a depth of water extraction of 80 or 100 cm. The crop with an 80-cm depth of water extraction was terminated in the simulations on Day 103.

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Fig. 3. Yield change in each year from a maize crop with an 80-cm depth of water extraction to a crop with 100- or 120-cm depth of water extraction plotted against yield simulated for the crop with an 80-cm depth of extraction.

Fig. 5. Yield change in each year from a maize crop with a relatively rapid leaf appearance rate to a crop with a relatively slow leaf appearance rate plotted against yield simulated for the crop with a relatively rapid leaf appearance rate.

not quite reached at this time, and subsequent precipitation increased the FTSW. The crop with a 100-cm depth of water extraction was simulated to progress normally to maturity and to eventually yield 840 g m2. An increased depth of soil water extraction increased crop yield in every year (Fig. 3). The greatest yield increase as a result of a greater depth of soil water extraction was in 1982. The increased access to water and a greater reservoir of plant available soil water increased the mean yield to 507 g m2 for the 100-cm depth, giving a mean yield increase of 98 g m2 compared with the 80-cm depth; for the 120-cm depth, the mean yield increased to 563 g m2154 g m2 greater than the 80-cm depth. The viability of maize for cropping in this environment was simulated to be markedly improved by selecting soils and cultivars that result in a greater depth of soil water extraction. An important consequence of the deeper rooting was that early crop termination was simulated to be either delayed or completely eliminated in a number of years. Out of the 11 yr in which early termination was simulated for the 80-cm rooting depth, termination was delayed with a 100-cm depth in 6 yr, and it was delayed with a 120-cm depth in 9 yr. The minimization of the catastrophic yield losses in those years with potentially early crop termination is likely to be of special economic benefit to growers.

Leaf Size
While a smaller leaf size resulted in a substantial yield advantage in 1982 by avoiding a fatal FTSW, this was a unique case among the 20 yr (Fig. 4). No other year offered a weather scenario that resulted in an increased yield over the yield achieved by the nominal maize simulations. The decreased light interception as a consequence of the smaller leaves resulted in decreased yields in other years so that the mean yield for the 20 yr was 39 g m2 less than the nominal simulations. Therefore, the smaller leaf size is not advantageous for increasing yields in this water-limited environment.

Rate of Leaf Appearance

Slowing the rate of leaf appearance, and thereby the rate of water use, resulted in the avoidance of a lethal FTSW in 2 yr (1974 and 1982). Consequently, large yield increases were simulated in these 2 yr (Fig. 5). The yields in many years also benefitted from the longer period of vegetative growth as a result of the slowed leaf appearance rate. The benefit of the extended growth period was greater for the higher yield levels. On the other hand, the slowed leaf appearance decreased yields in 5 yr. In 2 yr, the lengthening of the growing season resulted in premature senescence. Overall, the mean simulated crop yield was increased to 541 g m2, which is an increase of 132 g m2 over the nominal case.

Carbon Dioxide Assimilation

Similar to the preceding simulations, which had a decreased soil water use rate, decreased RUE led to sufficient soil water conservation to avoid a lethal FTSW in 1982, resulting in a substantial yield increase (Fig. 6). In many years, however, the slower growth rate resulted in decreased yields. This was particularly true in years with higher yields and high amounts of precipitation. Overall, the mean yield simulated for the decreased RUE was 421 g m2, which was slightly greater than the nominal case.

Fig. 4. Yield change in each year from a maize crop with a leaf area of the largest leaf (AMAX) of 750 cm2 to a crop with a largest leaf of only 500 cm2 plotted against yield simulated for the crop with a largest leaf of 750 cm2.

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Fig. 6. Yield change in each year from a maize crop with a radiation use efficiency (RUE) of 1.6 g MJ1 to a crop with an RUE of 1.25 g MJ1 plotted against yield simulated for the crop with an RUE of 1.6 g MJ1.

Fig. 8. Yield change in each year from a maize crop with a maizelike linear increase in harvest index (DHI) of 0.015 HI d1 to a crop with a greater DHI of 0.0185 HI d1 plotted against yield simulated for the crop with a maize-like DHI.

Early Stomata Closure

A decrease in RUE and water use by an early closure of stomata as the soil dried resulted in the avoidance of the lethal FTSW and an increased yield (Fig. 7a) in 1982 and two additional years. The remaining 17 yr showed surprisingly little response in the final yield to this early down regulation with a mean yield equal to 445 g m2. Because there appeared to be little negative consequence in introducing this trait, early stomata closure might be a potential plant trait for modification in environments where the development of a lethal FTSW is common.

Delayed Stomata Closure

Not surprisingly, delaying the down regulation in plant activity, which is claimed to be a major advantage of osmotic adjustment, resulted in a rapid use of the soil water and increased the likelihood of the crop being subjected to a lethal FTSW. With this change, the crop escaped the development of a lethal FTSW in this environment for only 4 out of the 20 yr. As a result, there was little or no increase in yields in any years, and in some years, yield was decreased (Fig. 7b). The mean simulated yield was actually decreased by 28 g m2 for delayed stomata closure compared with the standard case. These results directly challenge the putative benefit of maintaining leaf turgor by osmotic adjustment under water-deficit conditions.

Grain Growth and Duration

An increase in the DHI allowed for a shortening of the length of the grain growth period by increasing the rate of grain growth such that the crop might avoid the severe stress that could develop late in the season. This approach failed, however, to eliminate the effects of water deficit on yield (Fig. 8). In several years, this alteration resulted in substantial yield decreases because the threshold HI of 0.2, at which stress-induced termination was reached, occurred at an earlier date. In all years, yield was decreased to some extent by the shortening of the crop growing duration so that less crop mass was accumulated. With a mean yield of only 368 g m2, this plant trait resulted in the lowest mean yield of any trait tested.

Combined Traits
The combined sorghum-like traits produced a yield benefit of about 400 g m2 in 2 yr (1974 and 1982), and a benefit of about 200 g m2 in another 2 yr (1977 and 1988). Small benefits of the sorghum-like traits were also simulated for all situations where the yield of the maize simulations gave yields 550 g m2 (Fig. 9). On the other hand, in those environments in which maize yields were simulated at 550 g m2, yields were decreased when sorghum-like traits were used in the simu-

Fig. 7. Yield change in each year from a maize crop with normal stomata closure with respect to the fraction of transpirable soil water (FTSW) to a crop with (A) an early closure of stomata and (B ) a late closure of stomata plotted against yield simulated for the crop with normal stomata closure.

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Fig. 9. Yield change in each year from a crop with maize-like traits to a crop with a combination of sorghum-like traits plotted against yield simulated for the maize crop.

lations. The net result for the conditions of these simulations was that shifting the maize crop to a sorghum-like crop increased the mean mean 20-yr yield to 474 g m2, which was an increase of 75 g m2. One of the main benefits of the combined sorghum-like traits is a somewhat stabilized yield level achieved by increasing yields in the poor years and decreasing yields in the good years. The benefit of the decreased rate of leaf appearance, in particular, was suppressed in the good years by the other sorghum-like traits.

DISCUSSION
This analysis of putative plant traits to increase crop yields under water-limited conditions was undertaken as an approach particularly relevant to growers for increasing water use efficiency. That is, growers are more likely to benefit from increased yields on a finite water resource than from simply increasing water use efficiency per se. Consequently, a number of putative traits for improving crop performance under water-limited conditions were tested in a simulation of maize growth and yield. Similar to previous analyses, these simulations demonstrated the advantage of increasing the depth at which crops extract soil water (Table 1). Yields in all years of the 20 yr of simulations were benefitted by an increased depth of water extraction (Fig. 3). It seems clear that efforts to increase the soil water extraction depth, particularly if a crop is currently limited to a depth as shallow as 80 cm, are likely to be rewarded with increasing crop yields. Traits that decreased the rate of soil water extraction did not, in general, offer much benefit in yield increase (Table 1). Decreased leaf size and RUE resulted in little or no yield gain. These traits offered a benefit only in one year (1982) where the slowed soil water extraction rate allowed the lethal water stress to be avoided. The results from a study of six cultivars of pigeonpea [Cajanus cajan (L.) Huth] with differing RUE under irrigated conditions similarly showed no significant correlation of grain yield under water-limited conditions with irrigated RUE (Nam et al., 1998). On the other hand, decreasing the water extraction rate by slowing the rate of leaf development resulted in yield increases both in those years vulnerable to lethal

soil water deficits and in many other years with more adequate water. However, yield was decreased in 5 yr as a result of slowed leaf development, showing a variable response in yield to changes in this trait. The simulated yield decreases are consistent with the results of Grumet et al. (1987) from field experiments with barley (Hordeum vulgare L.) in which a comparison of isopopulations with differing rates of leaf development resulted in lower grain yields for the lines with slow leaf development. Altered stomata closure relative to the soil water content offered few benefits (Table 1). While early stomata closure allowed the avoidance of the lethal soil water content in the simulations for 1982, this trait had little influence on yield in most years (Fig. 7a). The recommendation for osmotic adjustment to delay stomata closure with soil drying proved generally to be deleterious (Fig. 7b). Any putative benefit of osmotic adjustment would likely have to influence crop survival during periods of exposure to potentially lethal periods of soil water deficit. The simulations that brought together the various sorghum-like traits in the maize model illustrated the benefit of these traits for dryland conditions. In all cases where the yield level calculated with the maize model was 550 g m2, crop yield was increased by adopting the sorghum-like traits. These results highlighted those traits of sorghum that make it the preferred crop for water-limited environments. In contrast, in those years when maize yields were simulated to be 550 g m2, crop yields were decreased by incorporating sorghumlike traits. These simulation results are consistent with the experimental comparison of maize and sorghum yields reported by Muchow (1989) for water-limited conditions in Katherine, Australia. In that study, maize yields were superior to sorghum yields when maize yields were at least 600 g m2. Sorghum yields were superior when maize yielded 600 g m2. Consequently, sorghum and sorghum-like traits appear appropriate only for very water-limited environments as a means to achieve greater yields, and thus, a greater yield for the fixed amount of available water. Overall, these simulations are encouraging in showing that crop traits can be adjusted to change yield under water-limited conditions. In particular, sorghum-like traits were generally advantageous in the simulated environments under low-yielding conditions (550 g m2), and maize-like traits were generally advantageous under high-yielding conditions (550 g m2). Further, the yield modifications directly resulted in shifts of the ratio between grain yield and evapotranspiration (Table 1), which is directly relevant to the concerns of growers for achieving the most efficient use of water. REFERENCES
Birch, C.J., P.S. Carberry, R.C. Muchow, R.L. McCown, and J.N.G. Hargreaves. 1990. Development and evaluation of a grain sorghum model based on CERES-Maize in a semi-arid tropical environment. Field Crops Res. 24:87104. Bolanos, J., G.O. Edmeades, and L. Martinez. 1993. Eight cycles of selection for drought tolerance in lowland tropical maize: III.

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