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Phyllochron Responds to Acclimation to Temperature and Irradiance in Maize

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Phyllochron Responds to Acclimation to Temperature and Irradiance in Maize 1 Phyllochron responds to acclimation to temperature and irradiance in maize. C. J. Birch1, J. Vos2, J. Kiniry3, H.J. Bos, 2 and A. Elings4 1. The University of Queensland, Gatton College, 4345, Queensland Australia, 2. Wageningen Agricultural University, Haarweg 333, Wageningen, The Netherlands. 3. United States Department of Agriculture, Grassland, Soil and Water Research Laboratory, 808 East Blackland Rd., Temple, Texas, 76502, USA 4. International Maize and Wheat Improvement Centre, Lisboa 27, Apdo Postal 6- 601, 06600 Mexico, D. F., Mexico Prepared for publication in: Field Crops Research Short Title: Phyllochron responds to acclimation and irradiance Corresponding Author: C. J. Birch, The University of Queensland, Gatton College, 4345, Queensland, Australia Phone 61 7 54601302 Fax 61 7 54601112 Email c. [email protected] 2 Abstract Crop models need accurate simulation of leaf canopy development. The thermal interval for leaf tip appearance (phyllochron) is critical for predicting the duration of vegetative development. The phyllochron in maize is shorter in temperate than in tropical and subtropical environments. As existing data has been evaluated in a narrow range of environments, the underlying mechanisms that affect phyllochron have not been adequately examined. The objectives of this study were to quantify the response of phyllochron to environmental variables and determine its stability across maize cultivars, to aid modelers in developing tools which accurately predict phenology. Maize was grown in field experiments at Wageningen, The Netherlands, Temple, Texas, USA, and three sites in Mexico, and in controlled environments at Wageningen. The experiment at Temple included grain sorghum and shading treatments to alter irradiance of the crop. Detailed data on leaf production and environmental conditions were collected. These data were combined with published data from field studies. Maize phyllochron acclimated to temperature and increased as mean daily temperature before tassel initiation increased from 12.5 to 25.5 oC, and increased in maize and sorghum in response to low irradiance. Temperature was the dominant influence, with phyllochron increasing by 1.7 oCd per oC increase in daily mean temperature, as daily mean temperature before tassel initiation increased from 12.5 to 25.5 oC, and declined or remained constant when mean daily temperature before tassel initiation exceeded 25.5 oC. Only small differences in phyllochron occurred among cultivars. Phyllochron increased by 2 to 4 oCd per MJ photosynthetically active radiation (PAR) as irradiance decreased from 9.6 to 1.1 MJ PAR m-2 d-1. Key words: acclimation, irradiance, leaf appearance, maize, modelling, phyllochron, sorghum, Sorghum bicolor, temperature, Zea mays 3 Introduction Crops develop and produce leaf canopies differently in cool temperate areas than in the tropics and subtropics. Phyllochron (thermal interval between the appearance of successive leaf tips) of maize in The Netherlands was lower (Struik 1983a,b, Hussen 1995) than in warmer temperate environments (Kiniry and Bonhomme 1991) and tropical and sub-tropical environments (Singh 1985, Bonhomme et al., 1991, Carberry and Abrecht 1991, Kiniry and Bonhomme 1991, Birch et al. 1998a). The phyllochron (oCd leaf-1) is fairly stable at a location while differing greatly among tropical, warm temperate and cool temperate latitudinal zones. Typically, a maize phyllochron is 30% greater in tropical than in temperate areas (Kiniry and Bonhomme 1991). These authors showed that the phyllochron was lower when mean minimum temperature of the four coolest nights in 30 days after emergence was less than about 8 oC than when above 12 oC. However, the relationship was not sufficiently robust for use in modelling. Accurate information on phyllochron is needed to predict crop development from leaf number. Models can use leaf number and a constant phyllochron to predict time of flowering as in CERES-Maize (Jones and Kiniry 1986), AUSIM-Maize (Carberry and Abrecht 1991), and modified models derived from them (Keating et al. 1991, Birch 1996). CORNF (Stapper and Arkin 1980) uses a similar approach, but with the appearance of ligules rather than leaf tips, while Muchow et al. (1990) use an exponential relationship that produces declining phyllochron as plants age. Sorghum models use a constant phyllochron (Rosenthal et al. 1989, Birch et al. 1990). Environmental modification of phyllochron causes errors in prediction by models that use a constant phyllochron, and could explain some of the errors in predicting silking date reported by Kiniry and Bonhomme (1991) and Birch et al. (1998a). AUSIM - Maize and the modified form proposed by Birch (1996) predicted silking too late with the data of Struik (1983 a, b), due to the high phyllochron used. Birch (1996) reported that the predicted silking date was sensitive to the value of phyllochron, further emphasising the need for accuracy in this parameter. Phyllochron varied with radiation intensity at controlled temperature (Gmelig-Meyling 1973, Struik 1983c). 4 Small variations in phyllochron with photoperiod have been reported (Warrington and Kanemasu 1983, Gmelig-Meyling 1973, Coligado and Brown, 1975), though no differences were found by Birch (1996). Genotypic differences might contribute to differences in phyllochron among sub- tropical, tropical, and temperate environments. Genotypic variation was small or non- existent for maize planted on the same date (Bonhomme et al. 1991, Birch et al. 1998a), but may be larger at high temperatures (Ellis et al. 1992). Birch et al. (1998a) found no differences in phyllochron among temperate and tropically-adapted maize hybrids grown at Gatton, southeastern Queensland, Australia. Similarly, Bonhomme et al. (1991) found only minor differences in phyllochron among such genotypes at Guadeloupe. Thus, differences in adaptation of genotypes did not explain the large differences in phyllochron observed across environments. Until factors controlling phyllochron have been identified, crop models need a method of adjusting phyllochron according to environmental conditions. The use of locality- specific values inherent in the models listed earlier is inappropriate, as it requires model calibration for each location. One possible approach to predicting phyllochron is a photothermal coefficient, based on the concept that leaf production depends on assimilate supply (which depends on irradiance), and assimilate consumption (which depends on temperature, expressed as thermal time). Such an approach was proposed for rice (Nix 1976) and pearl millet (Ong and Squire 1984). Another option is the acclimation hypothesis proposed for barley and wheat by Kirby (1995). This hypothesis argues that rate of leaf appearance throughout development of wheat and barley depended on the temperature conditions soon after emergence. Plants grown under cool temperatures early produced leaves more quickly per day and per degree day than plants grown at warmer temperatures. More recently, Ntiamoah and Brule-Babel (1997) have reported longer wheat phyllochron in warm conditions. Such acclimation in maize has not been described. 5 Thus, two issues important for understanding and predicting phyllochron are the potential use of a photothermal quotient, and the influence of low temperature acclimation. Consequently a series of controlled environment and field studies were done in The Netherlands and Texas, USA, comparing the results with data from previous studies over diverse environments. The objectives were to (i) present original data on the effect of temperature, genotype and irradiance on phyllochron of maize grown in controlled environments and the field at Wageningen, The Netherlands, Temple, Texas, USA, and three sites in Mexico (ii) use the data from these experiments and data from literature sources to test the hypotheses that: (a) phyllochron is determined by the ratio of daily photosynthetically active radiation and daily thermal time; (b) phyllochron is constant across genotypes within environments; and (c) phyllochron is related to mean daily temperature during early plant growth (acclimation hypothesis of Kirby 1995). Information of the effect of irradiance on phyllochron in grain sorghum was gathered at the Texas site. Materials and Methods Field experiment, 1997, Wageningen The experiment was located at Wageningen Agricultural University, Latitude 51o 58'N, Longitude 5o 40'E. Three cultivars of maize, viz. LG22.42 (producer Limagrain, maturity rating 6, the latest maturity of the three, Anonymous 1997), LG11 (Limagrain, maturity rating 7.5, Anonymous 1997) and Lincoln (Van der Have, maturity rating 9, Anonymous 1997) were planted on 18 April, 6 May, 23 May, 3 June and 24 June 1997 in a split-plot design. Sowing dates were the main plots and cultivars sub-plots, in three replicates. Sub-plots were 9m x 3 m, with 4 rows 0.75 m apart. A fourth cultivar, Hycorn 42, of Australian origin was added on 3 and 24 June 1997, to extend the range of adaptation of cultivars. Fertilisers and water were applied 6 at rates to ensure non-limiting conditions of nutrient and water supply during crop growth. Pests were rigorously controlled. Phosphorus and potassium was applied before sowing. Nitrogen was applied at 30 kg ha-1 before sowing and 100 kg ha-1 as calcium ammonium nitrate 3 and 6 weeks after emergence. Plant population was 10 m-2. Maximum and minimum temperatures 1.50 m above ground and 0.05 m below ground, and total solar radiation were recorded nearby
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