Agricultural and Forest Meteorology 150 (2010) 606–613 Contents lists available at ScienceDirect Agricultural and Forest Meteorology journal homepage: www.elsevier.com/locate/agrformet Growth and development of maize (Zea mays L.) in response to different field water management practices: Resource capture and use efficiency Liu Yi a,b,c,1, Yang Shenjiao a, Li Shiqing a,c,*, Chen Xinping a,c, Chen Fang b a State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau, Northwest A & F University, Yangling 712100, China Wuhan Botanical Garden, Chinese Academy of Sciences, Wuhan 430074, China c Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resource, Yangling 712100, China b A R T I C L E I N F O A B S T R A C T Article history: Received 23 June 2009 Received in revised form 20 January 2010 Accepted 1 February 2010 Soil–water supply is the main factor limiting crop production across the Loess Plateau in China. A twoyear field experiment was conducted to evaluate three possible water management practices – film mulching (FM), supplementary irrigation (SI) and rain-fed (RF, control) – in terms of resource capture and use efficiency in maize (Zea mays L.) in this area. The cumulative intercepted photosynthetically active radiation (PARi), air thermal time (TTair), soil thermal time (TTsoil) and evapotranspiration (ET) were monitored during both crop growing seasons, and the effects of the three treatments on the growth dynamics and grain yield (GY) of the maize crops were compared. The results showed that the FM treatment significantly accelerated development of the crop plants, and the SI treatment induced more rapid development in the vegetative stage than the RF treatment. Both FM and SI treatments markedly increased the shoot dry matter (DM) and GY (p < 0.05). The cumulative PARi, TTair and TTsoil during the reproductive stage were all significantly increased by both the FM and SI treatments relative to the RF treatment (p < 0.05), correlating well with observed increases in DM and GY. Both the FM and SI treatments also resulted in significantly higher (p < 0.05) radiation use efficiency, and the FM treatment significantly increased the water use efficiency, by 23–25%, in both years (p < 0.05). The results show that the tested water management practices have significant effects on soil moisture and thermal conditions, and hence the rates of growth and development of maize, in fields on the Loess Plateau, China. ß 2010 Elsevier B.V. All rights reserved. Keywords: Loess plateau Water management practice Maize Radiation use efficiency Water use efficiency 1. Introduction The productivity of crops is directly related to their capture of resources, such as water and light, and the efficiency with which they convert these physical resources into biological materials. These relationships allow growth efficiency to be quantified in terms of either a ‘solar engine’ or a ‘water engine’ model by relating their dry matter (DM) production to the amount of radiation captured or the amount of water transpired, respectively (AzamAli et al., 1994; Yang et al., 2004). Radiation use efficiency (RUE) may be defined as the amount of dry biomass (above ground or total dry matter) produced per unit intercepted solar radiation. RUE has often been considered to be constant for a given crop species in productivity modeling studies * Corresponding author at: Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resource, Yangling 712100, China. Tel.: +86 29 87016171; fax: +86 29 87016171. E-mail addresses: [email protected] (L. Yi), [email protected] (L. Shiqing). 1 Tel.: +86 15926293185. 0168-1923/$ – see front matter ß 2010 Elsevier B.V. All rights reserved. doi:10.1016/j.agrformet.2010.02.003 (Sinclair, 1986; Yang et al., 2004), but it may vary during plant growing seasons (Lecoeur and Ney, 2003; Werker and Jaggard, 1998) and is heavily dependent on climatic conditions, i.e., abiotic stresses can significantly reduce both the interception of radiation and RUE. For instance, under drought stress, plants may need to modify their water extraction patterns from the soil and minimize water losses by closing their stomata, reducing their leaf area expansion and even, in extreme cases, losing leaf area through abscission and/or senescence to survive and reproduce (Craufurd and Wheeler, 1999; Mwale et al., 2007a). Thus, reductions in canopy photosynthetic capacity due to restriction of CO2 diffusion into the leaves following stomata closure, reductions in intercepted photosynthetically active radiation (PARi) and losses of leaf area may all cause reductions in RUE, dry matter production and (hence) productivity loses (Collinson et al., 1996; Craufurd and Wheeler, 1999; Mwale et al., 2007b). Soil–water depletion and plant water use efficiency (WUE) are critical factors affecting agricultural productivity in arid and semiarid areas around the world. Hence, various soil and crop management practices have been developed to increase crop yields (Huang et al., 2005; Fang et al., in press), notably plastic or L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 straw mulching, which may efficiently improve the microclimate and crop growth conditions (Albright et al., 1989) by promoting plant transpiration at the expense of evaporation from the soil (Raeini-Sarjaz and Barthakur, 1997; Wang et al., 2009). Thus, both crop yields and WUE have often been reported to be increased by mulching treatments (Li et al., 2001; Li and Gong, 2002). Irrigation may also have beneficial effects on plant water relations and yields, but Kang et al. (2002) found that grain yield (GY) and WUE responses to irrigation varied considerably with differences in soil–water contents and irrigation schedules. Further, Wang et al. (2002) and Fang et al. (in press) showed that scheduled irrigation based on crop responses to water stress at different development stages can improve WUE, but Olesen et al. (2000) found that although irrigation increased yields, there were no significant differences in WUE and harvest index in wheat subjected to three different irrigation strategies, since the increases were almost solely due to increased transpiration. In addition, excessive irrigation can reduce crop WUE (Jin et al., 1999). The Loess Plateau, the cradle of traditional Chinese culture, covers an area of 623,800 km2 in northwest China and has a population of about 90 millions. Its climate is mostly semi-arid, with annual precipitation ranging from 150–300 mm in the north to 500–700 mm in the south (Li and Xiao, 1992). Most of the annual precipitation (50–60%) falls as rain from June to September. Groundwater resources are sparse and deep, so most of the agriculture on the Loess Plateau is dryland farming, relying solely on rainfall (Li and Xiao, 1992). Soil moisture supply is the main factor limiting crop production in the area, while the amount of radiation received is usually non-limiting (Li et al., 2004; Liang et al., 2006; Liu and Zhang, 2007). Maize (Zea mays L.) is one of the major crops on the Plateau, accounting for 27.3% of the total agricultural area (Xue et al., 2008). Quantifying the relationships between RUE, WUE, biomass accumulation and yields would be of great practical value to facilitate elucidation of crops’ physiological responses, and crop modeling, breeding and improvement in the area (Kiniry et al., 1998; Awal et al., 2006). The objectives of this study were: (i) to quantify the amount of solar radiation and soil–water captured by maize crops on the 607 Loess Plateau during the growing season, (ii) to analyze the efficiency with which the captured resources are used to produce biomass and grain yield, and (iii) to determine the effects of selected soil–water management practices on resource capture and use efficiency. 2. Materials and methods 2.1. Site description The Changwu experimental station (35.28N, 107.88E, ca. 1200 m above sea level) is situated on loessial tableland where the loess is more than 100 m thick. The soils are Cumuli-Ustic Isohumosols, according to the Chinese Soil Taxonomy (Gong et al., 2007), and contain 37% clay, 59% silt and 4% sand with a pH of 8.4 and a bulk density of 1.3 g cm3. The organic matter, total nitrogen, available phosphorus, available potassium and inorganic nitrogen contents are 11.8 g kg1, 0.87 g kg1, 14.4 mg kg1, 144.6 mg kg1 and 3.15 mg kg1, respectively. The average annual precipitation from 1990 to 2008 was 578 69 mm, with 55% falling between July and September, and the annual average temperature 9.2 2.3 8C. One crop a year of wheat or maize is produced by rain-fed agriculture across most of the region. The study presented here was conducted in the years 2007 and 2008. Daily meteorological information and water supplies (including rainfall and irrigation) during each of the growing seasons are presented in Fig. 1. 2.2. Experimental design and treatments Three water management treatments – rain-fed (RF), film mulching (FM), and supplementary irrigation (SI) – were applied to maize plots at the experimental station. The soil–water supply for both the RF and FM treatments was solely dependent on natural rainfall, while in the SI treatment soil moisture was maintained at 70–85% of the field water capacity (FWC) in 2007 and 70–80% of the FWC in 2008 by increasing the soil–water content to the upper limit when it dropped below the lower limits via furrow irrigation with tap water. The crop was irrigated in this manner five times in Fig. 1. Daily meteorological and water supply data during the monitored growth seasons in 2007 and 2008. 608 L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 2007 (May 8, May 20, June 14, July 14 and August 15) and four times in 2008 (May 22, June 5, July 7 and August 4), with irrigation depths on each occasion of 33.7 mm and 25.6 mm, determined using water meters in 2007 and 2008, respectively. The treatments were applied to 50.7 m2 (7.8 m 6.5 m) plots arranged in a completely randomized block design, with four replicates per treatment. In each plot, maize was grown by Ridge cultivation (a common maize cultivation practice across the Loess Plateau) in both years. Base fertilizer – consisting of 110 kg N ha1 in the form of urea (N 46%), 50 kg P ha1 in the form of calcium superphosphate (P2O5 12%), and 100 kg K ha1 in the form of potassium sulfate (K2O 45%) – was broadcast over the soil, which was then turned over by plowing to transfer the fertilizer to the subsurface. Ridges (ca. 0.45 m wide at the top with 0.15 m furrows at the base) were then constructed by banking up soil from two sides to a height of 0.1 m from the base. In the plots mulched with plastic, plastic film (0.7 m wide and 0.005 mm thick) was used to cover the soil surface of the ridges, but not the furrows, and the edges were secured under the soil in the bottom of the furrows. Maize (Zea mays L. pioneer 335) was sown in 5 cm deep holes spaced 0.2 m apart, using a manually powered hole-drilling machine, along the midline on the top of each ridge on April 20 and April 18 in 2007 and 2008, respectively. Before backfilling, 300 ml water was poured into each seed hole to encourage seedling emergence. Additional nitrogen, in the form of urea, was applied using a hole-sowing machine in the furrows at the jointing and tasseling stages, at rates of 80 kg N ha1 and 90 kg N ha1, respectively, following a nutrient management plan aimed to achieve a final yield of 14 t ha1. Maize cobs were harvested gradually, when they were ripe, from 28 August to 13 September in 2007 and from 8 to 20 September in 2008. Weeds were controlled manually as required during each crop growth season. 2.3. Sampling and measurements The sampled and measured maize plants and soil measurement sites were all at least 1 m from the field edge to minimize edge effects. The same, established sampling and measurement procedures were applied in both study years, as described below. 2.3.1. Maize development stage evaluation A standardized maize development stage system was used to identify plant development stages (Ritchie et al., 1992), and the date was recorded at which 50% or more of the maize plants in each plot reached the following vegetative (VS) and reproductive (RS) stages: planting time (PT), emergence stage (VE), tasseling stage (VT), silking stage (R1) and physiological maturity stage (R6). shape factor, k, empirically determined to be 0.75 for maize (McKee, 1964). Leaf Area Index (LAI) values for each plot were then calculated by multiplying the leaf area values by the plant density (85,000 plants ha1), i.e., LAI = leaf area (m2 plant1) 85,000 (plants ha1)/10,000 (m2 ha1). 2.3.4. Soil moisture and soil temperature determination The soil–water content was measured gravimetrically at 20 cm intervals within the 0–200 cm profile in each plot at the PT, V6, V12, R1, R3, R4 (only 2008) and R6, crop growth stages. The soil moisture in the 0–200 cm profile of each plot was then calculated by summing the moisture in all of its sampled layers. The soil temperature (0–5 cm) was recorded daily at 8.00 h and 14.00 h (the coolest and warmest times of the day, respectively) and averaged to obtain indications of the mean daytime soil temperature. 2.4. Data calculation 2.4.1. Intercepted photosynthetically active radiation (PARi) and RUE It was assumed that 50% of the total incident solar radiation (R) is photosynthetically active radiation (PAR), and the amount of PAR intercepted by the plant canopy (PARi) was computed using the following exponential function (Yang et al., 2004): X 0:5Rð1 ekLAI Þ PARi ¼ where R is the incoming total solar radiation (MJ m2 d1); k is the light extinction coefficient, which equals 0.65 for maize, according to Monteith (1969); and LAI is the leaf area index (m2 leaf m2 ground). RUE (RUEGY and RUEDM) was calculated as GY or DM in g m2 divided by the total amount of intercepted PAR (PARi) in MJ m2. 2.4.2. Evapotranspiration (ET) and WUE ET was determined by the following formula (Zhang et al., 1999, 2005b): ET ¼ DW þ P þ I where DW is the change in soil–water storage (mm) between planting and harvesting, P is the precipitation (mm) during the crop growing season and I is the total irrigation water quota (mm). The surface runoff and deep drainage are usually neglected in studies such as this in this area (Kang et al., 2002). WUE (WUEGY and WUEDM) was calculated as GY or DM in kg m2 divided by the total water use (i.e., ET) in mm. 2.4.3. Air thermal time (TTair) and soil thermal time (TTsoil) Cumulative air thermal time (TTair) (8C d) values were calculated using the base temperature (Tbase) of 10 8C (Miedema, 1982) and the equation (Tan et al., 2000a,b): X TT ¼ ðT mean T base Þ 2.3.2. Plant biomass sampling Three adjacent plants (at least 1 m from plot edges and 0.5 m from previous sample sites) in a row were sampled randomly from each plot, by cutting off the shoot at the first nodule on the stem, at the 6th leaf stage (V6), 12th leaf stage (V12), silking stage (R1), milk stage (R3), dough stage (R4, only 2008) and physiological maturity stage (R6). The harvested shoots were killed by heating them at 105 8C for 30 min, weighed after oven-drying for 24 h at 65–75 8C, and the total above ground biomass in each plot was expressed in terms of kg dry matter/ha. where Tmean is the daily mean atmospheric temperature, which was averaged from hourly measurements of air temperatures throughout each day. All Tmean < Tbase values were considered to be effectively equal to 0 8C (Arnold, 1974). Similarly, soil thermal time (TTsoil) was calculated using soil temperature data and Tbase = 10 8C. 2.3.3. Leaf area index (LAI) measurement The area of each of the fresh leaves of the sampled plants was determined immediately after harvesting them, by multiplying their manually measured length and maximal width with a The effects of the treatments on the measured parameters were evaluated by one-way ANOVA. When F-values were significant, the least significant difference (LSD) test was used for comparing between-treatment differences in means according to Duncan’s 2.5. Statistical analyses L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 new multiple range tests. In all cases differences were deemed to be significant if p <0.05. 3. Results Table 1 Averaged soil temperature and water storage over the growing season under the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) treatments in 2007 and 2008. Treatments 3.1. Soil moisture and thermal conditions FM significantly increased the average soil temperature, by 1.9–2.9 8C, in both years compared to the SI and RF treatments (Table 1). This may have been because solar energy passed through the mulch and heated up the air and soil beneath the mulch, then the heat was trapped by a ‘‘greenhouse effect’’ (Li et al., 1999; Zhou et al., 2009). SI enhanced the soil–water storage in the 0–200 cm profile compared to the FM and RF treatments (significantly in the 2008 growing season), but there was no significant difference in soil–water storage between nonirrigated treatments. 3.2. Crop development stages There were significant between-treatment variations in growth and development of the crop in both years (Fig. 2). Development was clearly promoted by the FM treatment, since the crops advanced to specific vegetative and reproductive stages earlier in the FM plots than in the SI and RF plots, e.g. seedling emergence was 3–4 d earlier, the crop reached the silking stage 5–12 d earlier; and the whole growth period from seedling emergence to physiological maturity was 15–17 d shorter. In the SI plots, irrigation induced rapid seedling and jointing growth in the VS, presumably because it maintained growth-promoting water Fig. 2. Durations of the vegetative stages (VS, from seedling emergence to silking) and reproductive stages (RS, from silking to physiological maturity) of the spring maize crops under rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) in 2007 and 2008. 609 RF FM SI Soil temperature (8C) Soil water storage (mm) 2007 2008 2007 2008 21.7 0.9 b 23.6 0.7 a 21.0 0.6 b 22.4 0.4 b 24.8 1.0 a 21.9 0.7 b 359.7 27.4 a 360.7 22.5 a 379.8 24.2 a 383.6 18.4 b 381.2 16.9 b 410.5 21.0 a Values are given as means standard error of means (n = 4). Values followed by different letters within a column are significantly different (p < 0.05). conditions (Çakir, 2004), so the silking stage was reached 6–10 d earlier than in RF plots, although the duration of the whole growth period was similar in both treatments. The duration of the VS and RS as proportions of the whole growth period also depended on the field water management practices (Fig. 2). The proportion for the vegetative stage was much higher in RF plots (56.9% in 2007 and 59.6% in 2008 on average) than in either FM (55.5% in 2007 and 53.8% in 2008, on average) or SI plots (51.1% in 2007 and 50.7% in 2008, on average). 3.3. Crop growth dynamics 3.3.1. Shoot biomass The shoot biomass was consistently significantly higher in the FM and SI plots than in the RF plots at each development stage (Fig. 3). This was presumably because film mulching increased the soil temperature by several degrees, thus enhancing growth of the maize during the early growth period, while the high soil– water content maintained by the SI treatment stimulated vegetative growth so the shoot DM content was much higher in SI plots at the physiological maturity stage (R6) than in the other plots. 3.3.2. Leaf area index (LAI) The LAI increased dramatically during the VS, peaked at the VT stage and then declined, indicating leaf senescence, in plots subjected to each of the treatments (Fig. 4), but there were substantial between-treatment differences in the LAI values. FM induced a rapid increase in LAI during the early VS (seedling stage), and hence an earlier peak and subsequent decline in LAI during the late development stages than the other treatments. The SI treatment resulted in a significantly higher maximum LAI than the other treatments, and prolonged the physiologically functional period in which the LAI was maintained at a high level from development stage R1 to R3, which should theoretically benefit Fig. 3. Time courses of changes in shoot dry matter (DM) in the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) plots in 2007 and 2008. Error bars are twice the standard error of the mean. V3–V8: from 3rd leaf stage to 8th leaf stage (seedling stage); V9–V18: from 9th leaf stage to 18th leaf stage (jointing stage); VT–R1: from tasseling stage to silking stage; R2–R4: from blister stage to dough stage; R5–R6: from dent stage to physiological maturity stage. 610 L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 Fig. 4. Time courses of changes in Leaf Area Index (LAI) in the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) plots in 2007 and 2008. Error bars are twice the standard error of the mean. V3–V8: from 3rd leaf stage to 8th leaf stage (seedling stage); V9–V18: from 9th leaf stage to 18th leaf stage (jointing stage); VT–R1: from tasseling stage to silking stage; R2–R4: from blister stage to dough stage; R5–R6: from dent stage to physiological maturity stage. Table 2 Grain yield (GY) and shoot dry matter (DM) under the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) treatments in 2007 and 2008. Treatments RF FM SI GY (t ha1) DM (t ha1) 2007 2008 2007 2008 12.2 1.1 b 14.6 0.6 a 15.2 0.8 a 11.4 1.4 b 14.2 1.0 a 15.6 1.8 a 19.3 2.4 b 24.1 1.3 a 24.7 1.8 a 20.5 3.8 b 24.0 2.1 a 26.5 3.4 a Values are given as means standard error of means (n = 4). Values followed by different letters within a column are significantly different (p < 0.05). assimilation, production and transportation in the plants (and hence the GY). 3.4. Grain yield (GY) and shoot dry matter (DM) GY and shoot DM values at harvest are shown in Table 2. Both FM and SI treatments significantly increased the maize GY compared to the RF treatment, by 19.7% and 24.6% in 2007, and by 24.6% and 36.8% in 2008, respectively. However, there was no significant difference in GY between the FM and SI treatments. Unsurprisingly, there were similar between-treatment differences in shoot DM at harvest. 3.5. Capture and distribution of the resources 3.5.1. Resource capture throughout the growing season The SI treatment resulted in significantly higher cumulative PARi and ET values throughout the whole growth season than the FM and RF treatments (Table 3), indicating that much more energy and water were utilized by the irrigated plants. In addition, the cumulative TTAir values were much lower in FM plots than in either SI or RF plots (probably due to the shortened growth season), but cumulative TTSoil values were significantly higher (which strongly compensated for the reduction in cumulative TTAir values). 3.5.2. Distribution of the resources in different development stages Unsurprisingly, the PARi was significantly higher during the reproductive stage, when the LAI was high, than during the vegetative stage for crops subjected to all treatments in both years. Both SI and FM significantly increased the PARi in RS compared to the RF treatment (Fig. 5A). SI, which significantly increased the transpirable soil moisture, promoted ET more than the FM and RF treatments during the whole growth season, but there was no significant difference in the ET between the latter two treatments (Fig. 5B). TTAir was significantly higher during the VS than the RS (because of the longer duration of the VS) in the RF plots, while it was similar in these two development stages in the SI and FM plots (Fig. 5C), and TTSoil was significantly lower during the RS in all plots, probably because of canopy shading, i.e. the reduction in the irradiation penetrating to the soil surface because of the high LAI (Fig. 5D). Both TTAir and TTSoil were significantly higher during the RS in the SI and FM plots than the RF plots. 3.6. Resource use efficiency (RUE and WUE) Similarly to the between-treatment differences in GY, grain yield RUE (RUEGY) and dry matter RUE (RUEDM) were significantly higher in both FM and SI plots than in RF plots, indicating that the irrigation and mulching treatments substantially improved the conversion of irradiation resources into economic crop yield components (Table 4). FM also significantly increased the grain yield WUE (WUEGY) compared to the RF treatment, by 23–25% in both years (Table 4), probably because of the restriction of water loss from evaporation, which makes no contribution to plant physiological performance, and the accompanying increase in plant transpiration. However, there was no consistent effect of irrigation on WUE during the two experimental seasons; SI increased the crop WUEGY (by 12%) in 2008 relative to the RF treatment, but significantly decreased it in 2007 (Table 4). Table 3 Whole season cumulative intercepted photosynthetically active radiation (PARi), evapotranspiration (ET), air thermal time (TTAir) and soil thermal time (TTSoil) under the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) treatments in 2007 and 2008. Treatments RF FM SI PARi (MJ m2) ET (mm) TTAir (8C) TTSoil (8C) 2007 2008 2007 2008 2007 2008 2007 2008 717 18 b 717 21 b 753 26 a 750 27 b 726 30 b 834 33 a 372 27 b 358 19 b 502 25 a 369 23 b 373 25 b 432 30 a 1358 30 a 1274 21 b 1357 27 a 1329 15 a 1303 14 b 1333 21 a 1564 65 b 1672 43 a 1462 74 c 1758 79 b 2018 54 a 1709 67 b Values are given as means standard error of means (n = 4). Values followed by different letters within a column are significantly different (p < 0.05). L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 611 Fig. 5. Accumulation of intercepted photosynthetically active radiation (PARi), evapotranspiration (ET), air thermal time (TTair) and soil thermal time (TTSoil) in spring maize in the vegetative stages (VS) and reproductive stages (RS) under rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) treatments in 2007 and 2008. Table 4 Whole season Radiation Use Efficiency (RUEGY and RUEDM) and Water Use Efficiency (WUE) (WUEGY and WUEDM) under the rain-fed (RF), film mulching (FM) and supplementary irrigation (SI) treatments in 2007 and 2008. Treatments RUEGY (g MJ1) RUEDM (g MJ1) 2007 2008 2007 RF FM SI 1.5 0.1 b 1.8 0.0 a 1.8 0.1 a 1.3 0.1 b 1.7 0.1 a 1.6 0.2 a 2.7 0.1 b 3.4 0.1 a 3.3 0.2 a WUEGY (kg ha1 mm1) WUEDM (kg ha1 mm1) 2008 2007 2008 2007 2008 2.7 0.1 b 3.3 0.2 a 3.2 0.2 a 28.5 1.5 b 35.6 1.2 a 26.4 2.0 c 27.0 2.7 b 33.1 2.6 a 31.4 4.5 ab 51.8 2.6 b 67.5 3.9 a 49.3 4.8 b 55.6 4.0 b 64.2 3.4 a 61.6 6.1 a Values are given as means standard error of means (n = 4). Values followed by different letters within a column are significantly different (p < 0.05). 4. Discussion The growth and development of a crop heavily depends on the species, variety (Ritchie et al., 1992), soil properties (Corwin et al., 2003), management practices (Mwale et al., 2007a,b), and meteorological conditions (Olasantan, 1999; Saini and Westgate, 2000). As a typical thermophilous crop, maize requires temperatures exceeding ca. 10 8C for its development and related physiological processes, such as canopy photosynthesis and root system activities (Miedema, 1982; Nguyen et al., 2009), and its biomass and GY are generally correlated to the whole season TT above this threshold (Ma et al., 2008). In our study, the TTair in the FM treatment was decreased, but the TTsoil was enhanced and the plant DM at maturity and the GY per 100 8C d were increased by 130–400 kg ha1 and 9–18% compared with 500 kg ha1 and 6.3%, respectively, in the study by Ma et al. (2008). The likely reason for this is that the FM treatment increased the heat resources available to the maize, which are crucial for initial dry matter production and for growth of maize in temperate climates (Xue et al., 2008; Ma et al., 2008). Both the SI and FM treatments improved the capture of sunlight, water or heat resources by the maize at the experimental site on the Loess Plateau, but by widely differing mechanisms. Generally, irrigation in drought-prone areas markedly promotes plant biomass accumulation because of the associated relief from drought stress (Çakir, 2004). In the present study, the SI treatment significantly advanced maize silking and markedly increased the durations of the reproductive stage and green LAI period compared to the RF treatment. It also improved the whole season PARi and ET, relative to the FM and RF treatments, in accordance with reports that improvements in water conditions during the VS (before silking) result in taller, more robust plants, larger leaf areas (NeSmith and Ritchie, 1992; Abrecht and Carberry, 1993), increases in vegetative dry matter (Claassen and Shaw, 1970a,b), and acceleration of leaf tip emergence, tassel emergence, silking, and onset of grain filling (NeSmith and Ritchie, 1992; Abrecht and Carberry, 1993). Maintaining a sufficient water supply during the RS (after silking) is also important since water deficits may extend the interval from silking to pollen shed (Herrero and Johnson, 1981) and curtail the grain filling period (Westgate, 1994). Thus, the SI treatment appears to have allowed the maize to exploit more sunlight, water and/or heat resources. In the FM treatment, the film mulch prevents water exchange between the soil and air, which in turn reduces the latent heat flux and the exchange of sensible heat between soil and air (Wang and Deng, 1991). At the seedling stage of crop development, the plant canopy was small, allowing most of the area covered by the plastic film to receive solar energy and the topsoil to warm up. Hence the mulched soil warmed more quickly than the unmulched soil during the day. Furthermore, it cooled more slowly at night because the plastic film and the water underneath the film reduces long wave radiation (Zhang et al., 2005a), even though the soil– water storage in FM plots was quite close to that in RF plots throughout the whole plant growth season (Table 1). The film also directly inhibits evaporation of water from the soil surface, while water moves from deeper soil layers to the topsoil by capillarity and vapor transfer, thereby keeping the topsoil–water content relatively stable (Wang et al., 1998; Li et al., 1999; Tian et al., 2003). 612 L. Yi et al. / Agricultural and Forest Meteorology 150 (2010) 606–613 Hence, the promotion of transpiration with little soil evaporation might have fostered biomass accumulation during early growth stages and accelerated plant development from seedling emergence to physiological maturity in the FM plots due to the stomata being more open than in the RF plots. Several previous studies have indicated that relationships between resource capture and crop productivity are affected by the temporal distribution of captured resources in different development stages (Mwale et al., 2007a,b). Notably, crop biomass production and GY appear to be most strongly related to resource capture during reproductive stages, since GY (especially) is directly associated with current rates of assimilation and translocation (Chen et al., 2003). Hence, grain yield can be dramatically reduced by sunlight, water or heat resources deficits during the RS (Bennett et al., 1989; Lindquist et al., 2005; Zhou et al. 2009). Results from the present study provide indirect support for this conclusion, since the cumulative PARi, TTair and TTsoil during the RS were all significantly increased by both the FM and SI treatments, accompanied by improvements in both total crop DM and GY. In other words, sunlight, water and heat resources deficits (Fig. 5) during the maize reproductive stage in the RF treatment relative to the FM and SI treatments might have caused inhibition of assimilation and photosynthate translocation, and hence decreased the total crop DM and GY. Furthermore, the SI treatment resulted in both the highest cumulative ET during the RS and the highest yields. However, crop productivity is not solely dependent on resource capture during the growing season, but also its use efficiency (Azam-Ali et al., 1994; Yang et al., 2004). Maize is characterized as having high RUE and WUE, and in the present study the crop RUEDM varied from 2.7 to 3.4 g MJ1 (depending on the water management treatment), within the range (2–4.3 g MJ1) reported in previous studies (Andrade et al., 1993; Yang et al., 2004); while the WUEGY ranged from 26 to 36 kg ha1 mm1 in the present study compared with 15–34 kg ha1 mm1 in previous studies (Kang et al., 2000; Deng et al., 2006; Fang et al., in press). SI improved both the PARi and RUE, thereby significantly increasing the GY, probably because it stimulated physiological processes after soil drying episodes, leading to compensation or overcompensation in plant growth and GY (Deng et al., 2006). However, the SI treatment did not result in consistent changes in WUE with respect to either total DM production or GY; the treatment deceased WUE in 2007, but increased it in 2008 when the amount of supplementary water was reduced. So, excessive irrigation may have reduced the crop WUE in 2007 (the total irrigation amounts were 168.5 mm and 102.4 mm in the years 2007 and 2008). Accordingly, Jin et al. (1999) found that increments in irrigation when soil moisture contents exceed a certain threshold may induce increasingly small yield increases, and even possibly reduce yields, and too much irrigation may merely enhance nonphysiologically active soil surface evaporation (Olesen et al., 2000), which may have contributed to the reduction of WUE in 2007. The FM treatment significantly improved both the RUE and WUE with respect to the total DM production and GY relative to the RF treatment, although the whole season cumulative PARi and ET were very similar in the two treatments. The FM treatment probably effectively restricted water loss via soil evaporation by creating an impermeable barrier, and instead increased the physiologically significant canopy transpiration (Raeini-Sarjaz and Barthakur, 1997; Wang et al., 2009), thereby enhancing plant physiological processes and (thus) plant productivity and GY (Li et al. 2001; Li and Gong, 2002) compared to the RF treatment. In contrast, large amounts of soil moisture were lost in the RF treatment through non-physiologically active soil surface evaporation, especially in early development stages when most of the soil surface was exposed to direct irradiation and a dry atmosphere. Hence, the plant growth activities were notably restricted by the ensuing water deficit, leading to reductions in both RUE and WUE. 5. Conclusion In conclusion, the capture of light, water and thermal resources (and their use efficiencies) responded sensitively to changes in soil moisture and thermal conditions under the tested water management practices in the maize fields on the Loess Plateau, China. Consequently, there were significant between-treatment differences in the chronology of crop growth and development, and final biomass and grain yield parameters. The results show that both SI and FM practices were beneficial for high yields of maize on the Loess Plateau. However, FM is likely to be generally preferable, given the shortage of available water resources in the area. Acknowledgements This work was supported by the National Basic Research Program of China (2009CB118604) and Natural Science Foundation of State Key Laboratory of Soil Erosion and Dryland Farming on the Loess Plateau (10502-Z04; 10501-247). References Abrecht, D.G., Carberry, P.S., 1993. 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