Intercepted Solar Radiation during Seed Filling Determines Sunflower Weight per Seed and Oil Concentration

bal,* Yorick Lavaud, Guillermo A. A. Dosio, Natalia G. Izquierdo, Luis A. N. Aguirreza lez. Fernando H. Andrade, and Leila M. Gonza ABSTRACT
A reduction in intercepted photosynthetically active radiation (PAR) during a short period of seed filling could affect weight per seed and oil concentration in sunflower (Helianthus annuus L.) depending when the reduction occurs. The main objectives of this work were (i) to determine the period of time during which intercepted PAR most influences final weight per seed and oil concentration, and (ii) to establish and validate relationships between intercepted PAR during such a period and oil yield components. Intercepted PAR was modified by shading (0, 50, or 80%) field grown crops during different periods of seed filling. Data from published and unpublished field experiments, in which intercepted PAR per plant was changed during the entire seed filling period by shading, thinning to reduce plant population, or both, were also used. All experiments were performed under adequate water and nutrient conditions. A reduction in intercepted PAR during 7 to 10 d of seed filling affected weight per seed and oil concentration. The developmental interval during which final weight per seed and oil concentration were best accounted for by accumulated intercepted PAR (r 2 0.805) began at 250C d and ended at 450C days after flowering (DAF). The established relationships were validated with independent data. The relationships obtained in this work help to explain the effects of environment and crop management (sowing date, location, year) on oil yield variation.

eight per seed and seed oil concentration in sunflower are determined during the seed filling period (from end of flowering to physiological maturity). Weight per seed is determined during the entire period bal et al., 1996). Seed oil concentration be(Aguirreza gins to increase rapidly a few DAF and continues to increase until a few days before physiological maturity, when it stabilizes (Goffner et al., 1988; Chervet and Vear, 1989; Connor and Hall, 1997). Changes in environmental conditions during the seed filling period could potentially affect both yield components. Relationships between yield components and environmental conditions have been established for several crops [e.g., Andrade et al., 2000, in maize (Zea mays L.); Wiegand and Cuellar, 1981, and Abbate et al., 1997, in wheat (Triticum aestivum L.)]. In sunflower crops grown under good moisture and nutritional conditions, seed number per unit land area was closely correlated to photothermal quotient [daily average solar radiation/ (daily average temperature base temperature)] from floret initiation to seed set stage (Cantagallo et al., 1997). Less information is available about the effect of environmental conditions on seed oil concentration (Hall, 2000)

and on weight per seed. Recently, weight per seed was closely related to intercepted PAR from the end of flowering (R6 46C d, base temperature 6C) to physiological maturity (R9, Schneiter and Miller, 1981) in two hybrids, one with high and the other with low seed oil concentration (Dosio et al., 2000). The variation in oil concentration in the hybrid with high seed oil concentration was also related to intercepted PAR during the same period (r 2 0.93). The duration of the R6 to R9 period can be highly variable as the time to physiological maturity varies among years in responses to environmental conditions. Therefore, relationships between intercepted PAR during R6 to R9 and weight per seed and oil concentration could then only be used a posteriori to describe rather than predict seed weight or oil concentration, or need to be coupled with more complicated models (i.e., Texier, 1992; Villalobos et al., 1996) to estimate the date of physiological maturity. To achieve a fast and simple prediction of both oil yield components, it could then be useful to establish relationships in which intercepted PAR is accumulated between fixed crop stages or fixed dates in time (e.g., Cantagallo et al., 1997). Assimilates used in grain filling mainly originate in current photosynthesis (Hall et al., 1988; Dosio et al., 2000). During the seed filling period, weight per seed shows a phase of fast increase during which sink demand for assimilates is strong. A phase of fast oil concentration increase has also been observed (Goffner et al., 1988). Therefore, a developmental interval during which final weight per seed and oil concentration are closely linked to intercepted PAR could exist. Determination of this critical developmental interval and of quantitative relationships between weight per seed and oil concentration and intercepted PAR per plant could be useful to improve sunflower crop management and modeling yield so that crops could be managed to maximize available environmental resources during such an interval. The objectives of this work were (i) to determine the period of time during which intercepted PAR per plant best accounts for final weight per seed and oil concentration, (ii) to establish relationships between intercepted PAR during such period and final weight per seed and oil concentration, and (iii) to validate these relationships using independent data. MATERIALS AND METHODS Cultural Details
Experiments were conducted at the INTA Balcarce Experimental Station, Argentina (3745 S, 5818 W). The soil was
Abbreviations: DAF, days after flowering; LDT, long-duration treatment; PAR, photosynthetically active radiation; SDT, short-duration treatment.

n Unidad Integrada Facultad de Ciencias Agrarias (UNMdP)Estacio Experimental Agropecuaria INTA Balcarce, C.C. 276, 7620 Balcarce, Argentina. Received 1 Nov. 2001. *Corresponding author (laguirre@ mdp.edu.ar). Published in Crop Sci. 43:152161 (2003).

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a Typic Argiudol. The hybrid used was Dekalb G-100 with an oil concentration from 480 to 540 g kg1 and a black hull. Experiments were performed under optimum water and nutrient conditions. Soil analysis indicated that neither N nor P fertilization were necessary as nutrient availability was adequate to obtain maximum yield of sunflower crops under nonlimiting water conditions (yield 5000 kg ha1; Sosa et al., 1999; Andrade et al., 2000). Soil water content was monitored every 5 to 7 d using a neutron probe (Troxler 4300, Troxler Electronic Laboratories, Inc., Research Triangle Park, NC). Irrigation application as needed maintained soil water above 40% of maximum available water in the first 0.60 m of the soil profile during the entire growing season. Pests and diseases were adequately controlled. Two experiments, hereafter referred to as Experiments SDT1 and SDT2 (Short-Duration Treatments 1 and 2) varied intercepted PAR per plant during short periods of the seed filling period. The experiments were designed as randomized complete blocks with three replications. Sowing dates were 14 Nov. 1995 (SDT1) and 2 Dec. 1999 (SDT2). The plots had four (SDT1) and six (SDT2) rows, 0.7 m apart and 6 m long. Emergence occurred 7 d after sowing in both experiments. Final plant density was 45 000 (SDT1) and 72 000 (SDT2) plants ha1. Flowering of a plant was defined by the appearance of stamens in all florets from the outer ring of the capitulum (R5.1 stage). Flowering of a plot was defined as when 95% of its plants had flowered, and occurred on 20 Jan. 1996 (1 d) and 4 Feb. 2000 (1 d) in SDT1 and SDT2, respectively. In SDT1, treatments included a control (no shading), shading from 255 (R6 46C d) to 332C d (Period 1), from 332 to 430C d (Period 2), from 430 to 620C d (Period 3), and during the entire seed filling (R6 46C d to R9, 620C DAF, Fig. 1a). The 0C day corresponded to the R5.1 stage and the base temperature was 6C. In SDT2, treatments consisted of 50 or 80% reduction of incident solar radiation during three different periods of the seed filling stage (Fig. 1d). Treatments were a control (no shading, Fig. 1d) and 50 or 80% shading from 190 (R6 10C d) to 316C d (Period 1), from 316 to 468C d (Period 2), and from 468C d to 620C d (Period 3). A long duration shading treatment was not applied in this experiment. In both experiments, plots were shaded with black, synthetic, and neutral mesh clothes stretched on a pipe structure.

to prevent absorption of solar and long wave radiation. Measurements began at R5.1 and were registered every 60 s. Data were averaged every hour, and recorded by a data logger (LI-1000, LI-COR). Physiological maturity (R9) was estimated visually from the hard yellow color of the capitulum back face and from the brown color of its bracts (Farizo et al., 1982). In the control treatment, three capitula per plot were harvested every 3 to 7 d during the seed filling stage to determine physiological maturity as the time when average weight per seed does not further increase (see Data Analysis). Both methods used to determine R9 in control treatments gave similar results. The R9 stage of the control treatment occurred 45 and 28 DAF in SDT1 and SDT2, respectively. Final yield components were determined on 10 capitula harvested at the last sampling date (53 and 45 DAF in SDT1 and SDT2, respectively). All capitula were sampled from the two central rows of each plot. In SDT1, achenes were separated manually into nonempty (kernel occupying at least 20% of total space in the pericarp) and empty. In SDT2, fruits were cleaned mechanically (manual ventilator). Nonempty fruits were counted, oven-dried (with air circulating at 60C) to constant weight, and weighed. Weight per achene was obtained by dividing the weight of all nonempty achenes by the number of nonempty achenes. Hereafter, seed will refer to the entire fruit or achene. Oil concentration was measured in duplicate samples by nuclear magnetic resonance (Analyzer Magnet Type 10, Newport Oxford Instruments, Buckinghamshire, UK) and averaged. Weight per seed and oil concentration were expressed on a dry weight basis.

Data Analysis
In each experiment, seed number, weight per seed, and oil concentration data were analyzed using an analysis of variance procedure (SAS Institute, 1988, PROCANOVA, General Linear Models Procedure). When statistical differences were detected in more than one experiment, only the highest P value is presented. Differences among treatment means were evaluated with the Tukey test (P 0.05). In the control treatment of SDT1 and SDT2, weight per seed and oil concentration during seed filling was a function of thermal time (base temperature 6C) with a linear increase followed by a plateau. Two-step conditional models were used to determine the rate and duration of the linear increase phase for weight per seed and oil concentration. Weight per seed (Ws) and oil concentration (O ) were in the first phase (TTAF c )

Measurements
Global daily incident radiation was measured with a pyranometer (LI-200SB, LI-COR, Lincoln, NE) located 400 m from the experiments. Daily incident PAR was calculated as 0.48 global daily incident radiation. The proportion of PAR intercepted by the crop at solar noon was determined according to Gallo and Daughtry (1986), as (1 Rb/Ro), where Rb is the radiation measured below the lowermost green leaf, and Ro is the radiation measured above the canopy. Rb and Ro were measured weekly at solar noon (1 h), with a line quantum sensor (LI-191SB, LI-COR). According to Charles-Edwards and Lawn (1984), the daily proportion of PAR intercepted was calculated as 2 proportion of PAR intercepted at noon/ (1 proportion of PAR intercepted at noon). In sunflower, this correction allowed a substantial improvement on the error arising from a single measure at noon (Trapani et al., 1992). Daily intercepted PAR was obtained from global daily incident radiation and the proportion of PAR intercepted by the crop at noon, as in Dosio et al. (2000). Air temperatures were measured with copper-constantan thermocouples (Thermoquar, Buenos Aires) at the capitulum level. Thermocouples were protected by homemade shields

Ws or O a b TTAF
In the second phase (TTAF c ),

[1] [2]

Ws or O a b c

where TTAF was thermal time after flowering, a is the intercept, b is the rate of the linear increase phase, and the constant c is the time when the maximum weight per seed or oil concentration was reached (Dosio, 1998). Regression analysis was used to determine the critical developmental interval during which sensitivity of weight per seed and oil concentration to intercepted radiation was highest. Final weight per seed and oil concentration for each experimental plot in SDT1 and SDT2 were linearly regressed against intercepted PAR accumulated during consecutive 100C d periods of the seed filling stage, regardless of whether the plots were shaded or unshaded during that period. The r 2 values of these relationships were recorded. For each variable, the critical interval was composed of those 100C d periods with r 2 0.37 (SDT1, n 15) or r 2 0.29 (SDT2, n

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Fig. 1. Chronogram of radiation treatments (a, d), weight per seed (Ws; b, e), and oil concentration (O; c, f) as functions of thermal time after flowering for the control treatment and Exp. SDT1 and SDT2. For a, d: each horizontal bar indicates the timing and duration of the shading period in thermal time with respect to flowering (0C d, base temperature 6C) for a single treatment. Gray bar: 50% shading intensity, black bar: 80% shading intensity, open bar: no shading (control). For b, c, e, f: lines represent control treatment data adjusted to linearplateau functions. Vertical bars represent the standard deviation of the mean of three replications.

21). These r 2 corresponded to a P value 0.01 (Little and Hills, 1990). Combined weight per seed and oil concentration data from SDT1 and SDT2 were related to PAR during the previously determined critical interval using linear or negative exponential functions. The function which best described the experimental data was chosen by comparing r 2 values of the linear regressions between observed and estimated data. Final weight per seed and oil concentration were also related to intercepted PAR during R5.1 to R9 and during R6 to R9. To investigate final weight per seed and oil concentration under a wider range of intercepted radiation during the deter-

mined critical interval, data from SDT1 and SDT2 were compared with other published and unpublished data obtained in six field experiments that include shading and thinning treatments during the whole seed filling period (LDT longduration treatments). These experiments are hereafter referred to as Experiments LDT1, LDT2a, LDT2b, LDT3, LDT4, and LDT5. Table 1 shows sowing and flowering dates, duration of the R5.1 to R9 period, plant density at flowering, applied treatments, and data source for these experiments. The experimental design and further details for LDT1 and LDT2a are described in Andrade and Ferreiro (1996) and for LDT2b, LDT3, and LDT4 in Dosio et al. (2000). Experimental

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Table 1. Sowing date, flowering date, duration of the period from flowering (R5.1 stage) to physiological maturity (R9 stage), plant density, and shading treatments for experiments including long duration radiation treatments. Percentage values indicate the shading and thinning intensity applied to each experiment.
Treatments Experiment LDT1 LDT2a LDT2b LDT3 LDT4 LDT5# Sowing date 14 22 30 16 14 17 Oct. 1992 Oct. 1993 Nov. 1993 Dec. 1994 Nov. 1995 Nov. 1997 Flowering date 7 Jan. 1993 11 Jan. 1994 8 Feb. 1994 17 Feb. 1995 19 Jan. 1996 1 Feb. 1998 R5.1R9 C d 729 605 408 546 639 415 Plant density plants 58 000 58 000 72 000 72 000 45 000 58 000 ha1 Shaded % 45 45 50 50 50 50 Thinned % 75 75 75 75 50 50 Sh-Th No No Yes Yes Yes No

Shading and thinning treatment. Andrade and Ferreiro, 1996. This treatment was not applied during the considered experiment. Dosio et al., 2000. # Izquierdo and Dosio, 1998, unpublished data.

Table 2. Cumulative intercepted photosynthetically active radiation (PAR) SD from the beginning of the first treatment period to physiological maturity, for short-duration treatments (SDTs) SDT1 and SDT2. Periods 1, 2, and 3 refer to the time of the seed filling stage at which shading was applied. The percentages (50 or 80%) refer to the shading level applied. In SDT1, Periods 1, 2, and 3 corresponded to 255 to 332, 332 to 430, and 430 to 620C days after flowering (DAF), respectively. In SDT2, Periods 1, 2, and 3 corresponded to 190 to 316, 316 to 468, and 468 to 620C DAF, respectively.
Treatments Exp. SDT1 SDT2 Control 60.8 2.9 32.7 5.3 Entire Filling, 50% 26.5 1.1 Period 1, 50% 52.7 2.4 20.8 1.8 Period 2, 50% Period 3, 50% Period 1, 80% 18.3 1.3 Period 2, 80% 18.0 1.6 Period 3, 80% 24.8 1.9

MJ plant1 51.3 2.5 42.7 1.4 22.7 1.5 30.2 1.9

Shaded during the entire seed filling stage. This treatment was not applied during the SDT1 experiment.

design and crop management for LDT5 (the only unpublished LDT data set) was similar to those described in Dosio et al. (2000). During the period from R6 to R9, temperature varied from 18.1 3.6C (LDT3) to 20.3 3.2C (LDT1) and incident PAR from 6.3 2.8 MJ m2 d1 (LDT3) to 11.6 1.7 MJ m2 d1 (LDT2a). In these experiments, treatments to vary intercepted PAR per plant were applied approximately at the end of flowering (R6) and ended at physiological maturity (R9). In all experiments, a shading (45 or 50% shading), a thinning, and a control treatment were included. The thinning treatments consisted of uniformly removing plants to 25 or 50% of the original plant density. In LDT2b, LDT3, and LDT4, a treatment combining shading (50%) and thinning (shadedthinned treatment) was also applied. A wide range of variation was obtained in cumulative PAR intercepted per plant during the seed filling stage (10.1 0.2 MJ plant1 for the shading treatment from LDT3 to 138.8 3.3 MJ plant1 for the thinned treatment from LDT2b). This range depended on the levels of shading and thinning, plant density, and the variability in incident PAR between experiments. Relationships between final weight per seed and final oil concentration and cumulative intercepted PAR per plant during the critical interval were established using data from SDT1, SDT2, LDT2a, LDT2b, LDT3, and LDT4. Prediction quality of the established relationships was validated by the comparison between the predicted values from these equations and independent experimental data from LDT1 and LDT5. The hypothesis of intercept 0 and slope 1 were tested (t test, P 0.05) for intercepts and slopes derived from regression equations between predicted and observed data. For each experiment, only the highest absolute t value is presented.

3.4C respectively) while daily mean incident radiation (PAR) showed an inverse trend (10.5 2.3 vs. 9.2 2.2 MJ m1 d1 respectively). Cumulative intercepted PAR per plant from the beginning of the first treatment period to R9 showed a wide range of variation in both experiments (Table 2). This range depended on shading levels, plant density, and duration of the period during which shading was applied.

Oil Yield Components

Weight per seed and oil concentration as a function of thermal time for the control treatment in SDT1 and SDT2 were well described by linear-plateau functions (r 2 0.96, Fig. 1). Shading treatments affected weight per seed and oil concentration in both experiments. In SDT1, weight per seed was decreased by the long shading treatment (P 0.02, Table 3) but not by the short shading treatments. In SDT2, weight per seed was reduced (P 0.0015) by the 50 and 80% short shading treatments during Periods 1 and 2 (Table 3). Oil concentration was affected by shading in both experiments (P 0.0001, Table 3). In SDT1, the lowest oil concentration was obtained when plants were shaded during the whole seed filling period. In both experiments, the strongest effect of shading during a short period was observed when intercepted radiation was reduced at the middle of seed filling. In SDT1, nonempty seed number per plant was reduced compared with the control only by the long shading treatment (P 0.007, Table 3). In SDT2, shading at an early stage of seed filling affected nonempty seed number per plant (P 0.0006, Table 3). Seed number

RESULTS Growth Conditions

Daily mean air temperature during seed filling was lower in SDT1 than in SDT2 (19.4 3.4 vs. 20.5

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Table 3. Oil yield components in response to shading treatments in hybrid G-100 for short-duration treatments (SDTs) SDT1 and SDT2. Periods 1, 2, and 3 refer to the portion of the seed filling stage in which shading was applied. The percentages (50 or 80%) refer to the shading level applied. In SDT1, Periods 1, 2, and 3 corresponded to 255 to 332, 332 to 430, and 430 to 620C days after flowering (DAF), respectively. In SDT2, Periods 1, 2, and 3 corresponded to 190 to 316, 316 to 468, and 468 to 620C DAF, respectively.
Filled seeds Treatments Control Whole filling, 50% Period 1, 50% Period 2, 50% Period 3, 50% Period 1, 80% Period 2, 80% Period 3, 80% SDT1 seed 1899a 1494b 1778ab 1932a 1879a plant1 1182a 884bc 1029ab 1087ab 747c 852bc 970abc 56.0a 45.4b 56.3a 48.5ab 50.9ab SDT2 Weight per seed SDT1 mg seed1 40.4a 32.6c 33.8bc 38.6ab 34.6bc 31.7c 36.4abc 513ab 455d 519a 476cd 492bc SDT2 Oil concentration SDT1 g kg1 472a 451ab 418bc 468a 407c 361d 442abc SDT2

For each variable and experiment, means followed by the same letter are not significantly different (Tukey, P 0.05). Shaded during the entire seed filling stage. This treatment was not applied during the considered experiment.

per plant, weight per seed, and oil concentration tended to be higher in SDT1 than in SDT2 (Table 3), in association with values reported for intercepted radiation (Table 2).

Relationships between Oil Yield Components and Intercepted PAR

Final weight per seed was closely related to intercepted PAR per plant accumulated from 300 to 500C DAF in SDT1 (r 2 0.370, P 0.01, Fig. 2a) and from 250 to 500C d in SDT2 (r 2 0.29, P 0.01). In this experiment, r 2 decreased after the 400 to 500C d period and R9 was attained at 450C d. On the basis of these data, the interval during which weight per seed was best accounted for by accumulated intercepted PAR began at 250C DAF and ended at 450C DAF. The interval during which final oil concentration was better accounted for by intercepted PAR was similar to the interval for weight per seed (Fig. 2b, 250450C d). Variability in accumulated intercepted PAR per plant for the 100C d periods within the selected interval were similar to accumulated intercepted PAR variability for the other 100C d periods (CV 25.6 vs. 26.0% and 34.6 vs. 38.8% for SDT1 and SDT2, respectively). This result makes it unlikely that the higher regression coefficients for these periods were due to greater variation in intercepted PAR. During the 250 to 450C d interval, 32.3 (SDT1) to 51.1% (SDT2) of final weight per seed, and 50.3 (SDT2) to 53.8% (SDT2) of final oil concentration were accumulated. Weight per seed was linearly and oil concentration curvilinearly related to cumulative intercepted radiation per plant during the 250 to 450C d interval after flowering (Fig. 3). These oil yield components were also related to cumulative incident radiation and to daily mean temperature during the same interval, but with r 2 values lower than those obtained with intercepted radiation (0.337 and 0.470 vs. 0.805 for weight per seed and 0.640 and 0.110 vs. 0.860 for oil concentration). An effect of radiation treatments other than via a reduction in intercepted radiation was therefore unlikely. Accumulating intercepted radiation during longer intervals (R6 to R9, and R5.1 to R9) slightly improved the relationship between intercepted radiation and weight per seed and

did not improve the relationship between intercepted radiation and oil concentration. Most of the variation in weight per seed from LDT2a, LDT2b, LDT3, and LDT4 was well accounted for by the relationship established with data from SDT1 and SDT2 (Fig. 4a, r 2 0.73, P 0.0001) provided that intercepted PAR during the 250 to 450C d interval was

Fig. 2. Coefficient of determination (r 2) corresponding to linear regressions calculated from (a) weight per seed (Ws) and intercepted photosynthetically active radiation (PAR), and (b) oil concentration (O ) and intercepted PAR, for different periods of 100C d during seed filling for Exp. SDT1 and SDT2 (0C d corresponded to flowering, base temperature 6C). Tick labels on horizontal axes of figures correspond to the beginning points of the 100C d periods. Horizontal lines indicate r 2 values corresponding to P 0.01 (r 2 0.37 and 0.29 for SDT1 and SDT2, respectively).

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lower than 37 MJ plant1. The data obtained in thinning treatments, in which intercepted PAR was high, were overestimated by this relationship (Fig. 4b) and were accounted for by plant density before thinning (r 2 0.802, P 0.0001, n 14). Data presented in Fig. 4 were well accounted for (r 2 0.781, P 0.0001, n 88, Fig. 4b) by a conditional model combining (i) the function established between weight per seed and intercepted radiation without data from thinned treatments (Fig. 3a) and, (ii) a maximum weight per seed for each initial plant density. Residuals from this conditional model were not related to mean temperature (P 0.0735) or seed number per plant (P 0.8844). When data from all experiments were combined, oil concentration was a function of cumulative intercepted PAR per plant during the 250 to 450C d interval after flowering (Fig. 5). The relationship showed an increased phase followed by a plateau. The function established using SDT1 and SDT2 data alone overestimated oil concentration for data obtained at high intercepted PAR (no data at high intercepted radiation were available from SDT1 and SDT2 experiments, Fig. 3b). A linearplateau function between oil concentration and intercepted radiation was adjusted using SDT1, SDT2, LDT2a, LDT2b, LDT3, and LDT4 data. This function accounted for 80.3% (P 0.001) of oil concentration variability. Oil concentration increased up to 26.3 MJ plant1. Beyond this value, there was no response to intercepted radiation and oil concentration stabilized at 500 g kg1. Residuals from the established relationship

were not related to seed number per plant, daily mean temperature during the 250 to 450C d interval after flowering, and plant density (P 0.08). A negative exponential function (r 2 0.793 vs. 0.803) did not improve the adjustment compared with the linear plateau function presented on Fig. 5. Final weight per seed depends not only on intercepted PAR during seed filling but also on seed number per plant. Weight per seed and oil concentration were better accounted for by intercepted PAR than by intercepted PAR per seed (r 2 0.781 vs. 0.310 for weight per seed and 0.793 vs. 0.483 for oil concentration). The range of variation across treatments and experiments for both variables was large (CV 65.2 and 48.4% for intercepted PAR and intercepted PAR per seed, respectively). Most of the variation in intercepted PAR per seed was due to differences among experiments. Increasing intercepted PAR during seed filling tends to enhance nonempty seed number per plant (Andrade and Ferreiro, 1996; Dosio et al., 2000; Izquierdo and Dosio, 2002, unpublished data). Weight per seed was not related to seed number in LDT3 (P 0.542) but a

Fig. 3. Final (a) weight per seed and (b) oil concentration as functions of cumulative intercepted radiation (PAR) from 250 to 450C d after flowering for Exp. SDT1 and SDT2. Data points represent values obtained in each experimental plot.

Fig. 4. Weight per seed (Ws) as a function of cumulative intercepted radiation (PAR) per plant from 250 to 450C day after flowering for Exp. SDT1, SDT2, LDT2a, LDT2b, LDT3, and LDT4. (a) The line represents the relationship established with intercepted radiation 37 MJ plant1 (equation in Fig. 3). (b) The regression represents the relationship established with intercepted radiation 37 MJ plant1 (equation in Fig. 3) and the horizontal lines represent the maximum Ws calculated for each plant density (Ws 81.64 4.75x ). Data points represent values obtained in each experimental plot.

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Fig. 5. Oil concentration (O ) as a function of cumulative intercepted radiation (PAR) per plant from 250 to 450C d after flowering for Exp. SDT1, SDT2, LDT2a, LDT2b, LDT3, and LDT4. Line represents the adjusted function. Data points represent values obtained in each experimental plot.

SDT2, LDT2a, LDT2b, LDT3, and LDT4 data were tested using independent data from LDT1 and LDT5 experiments. Weight per seed and oil concentration obtained in LDT1 and LDT5 experiments were well predicted by the relationships established by using data from the other experiments (Fig. 6). Regressions between observed and predicted data in all cases gave intercepts and slopes not statistically different from 0 and 1 respectively (t 2.09 and 1.76 for LDT1 and LDT5, respectively). The greatest prediction error was observed for weight per seed in LDT5 (Fig. 6c). Most of this prediction error was explained by underestimation in the shaded treatment as observed vs. predicted data from the thinning and control treatments were close to the 1:1 line.

DISCUSSION
A reduction in incident and intercepted solar radiation for a short period during the seed filling stage affected weight per seed and oil concentration. The reduction obtained in this work was in some cases lower than the reduction produced by a cloudy period of 10 d in the region in which experiments were performed (Gon lez, 2001, unpublished data). Our results could help za to explain the variability in weight per seed and oil concentration among years within a region for crops growing under adequate moisture and mineral conditions and relatively free of pests and diseases. Intercepted PAR from 250 to 450C DAF accounted for most of the variability in weight per seed and oil

positive relationship was found between the two variables in the other experiments (P 0.027). When analyzed for each experiment, relationships between oil concentration and seed number per plant showed a similar trend to those observed between weight per seed and seed number.

Validation
Prediction quality of the relationships between weight per seed and intercepted PAR and between oil concentration and intercepted PAR established using SDT1,

Fig. 6. Observed vs. predicted data of weight per seed (Ws) and oil concentration (O ) for Exp. LDT1 (a and b) and LDT5 (c and d). Predicted values were calculated from the functions given in Fig. 4b and 5. Solid lines are the y x lines and dashed lines show the linear regressions between observed and predicted values.

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concentration observed in the experiments in which intercepted PAR was reduced for a short period during the seed filling stage. Considering longer periods to accumulate the intercepted PAR (i.e., the whole seed filling) only slightly improved prediction of those yield components. It is noteworthy that intercepted radiation accumulated during 100C d periods placed within the critical 250 to 450C d interval accounted for a high proportion of the variability in both yield components. However, the proportion of weight and oil accumulated during the 250 to 450C d critical interval was 53.8%. These results suggest that intercepted radiation influenced both variables before the effect occurred. Such delayed effects, often called memory effects, have been reported for several plant processes such as root elonga bal et al., 1994), oleic acid percentage, tion (Aguirreza (Izquierdo et al., 2002), pollen production and viability (Chimenti et al., 2000), and potential weight per seed (Connor and Hall, 1997). The linear function obtained in the experiments with SDTs accounted for most of the variation in weight per seed obtained in experiments in which intercepted PAR was modified during the whole seed filling stage in different years and for different sowing dates. Nevertheless, weight per seed of thinned treatments did not increase when intercepted radiation accumulated during the critical period was 37 MJ m2. Since crops were cultivated under good moisture and mineral nutrition, weight per seed in thinning treatments could be considered as a maximum for these growth conditions. This suggests that the physical or chemical capacity to accumulate dry matter in seed tissues was saturated in plants intercepting high quantities of PAR and that this capacity was determined prior to the time at which the plots were thinned (end of flowering). Interestingly, weight per seed in thinning treatments was higher at lower initial plant density. These results suggest that maximum weight per seed depends on the resource availability previous to treatment application and that this resource level would determine anatomical seed features that define maximum capacity to accumulate dry matter in seed tissues. Our results agree with those found in other species and other environmental conditions {i.e., Munier-Jolain et al., 1996, in soybean [Glycine max (L.) Mer.]}. They also support the idea that stresses expressed before rapid seed filling could reduce the weight per seed at harvest (Connor and Hall, 1997). Potential weight per seed seems to be determined before the beginning of seed filling, but an increase in intercepted radiation during 250 to 450C DAF increased final weight per seed in nonthinned treatments. Increasing leaf area index during this period could help to obtain a higher weight per seed. This index is often lower than the value needed to obtain 95% interception bal of the incident solar radiation (2.5 to 3, Aguirreza et al., 1996). A higher leaf area index during this critical period could be attained through genetic improvement by selecting for longer leaf area duration of the lower leaf strata. More than 78% of weight per seed and oil concentration variabilities were accounted for by intercepted radi-

ation from 250 to 450C DAF, despite variations among treatments or experiments in plant density, temperature, incident solar radiation, and seed number per plant. This confirms the significant role of intercepted radiation during the seed filling stage in determining weight per seed and oil concentration (Dosio et al., 2000). In this work, we have determined the period of the seed filling in which these two oil yield components show the highest sensitivity to changes in this environmental factor. Moreover, in the relationships established here, the period during which intercepted radiation was accumulated to estimate weight per seed and oil concentration began and ended at fixed thermal times after flowering. This is an advantage for the use of these relationships in simulation tools, compared with relationships presented by Dosio et al. (2000). Some experimental work is still needed to better know the validity domain of the relationships in this paper. First, we did not study whether relationships could change under extreme values of seed number. It could be possible that our relationships do not perform well as prediction tools under poor seed set and good environmental conditions during seed filling. Although the range of source-sinks relationships investigated in this work was wide, relationships taking into account the source (intercepted radiation) effect alone better accounted for weight per seed and oil concentration than those taking into account the effect of source and sink (seed number). For this reason, we kept intercepted radiation as the explanatory variable in our relationships. Second, the relationships presented here should be tested in a wider range of temperatures to verify their validity. Temperature was not directly considered in our relationships as no effect of temperature was detected. Nevertheless, the range of average daily temperatures in our experiments (2.4C) largely covered the daily mean differences in air temperature between January, February, and March (2.3C for 19701995), the months during which seed filling could take place in our region. This range is within the range of temperature variation found during seed filling for normal sowing dates in most of the regions in the world in which sunflower is grown. This lack of temperature effect could probably be explained by the range in average temperatures in our experiments (18.120.5C), which was close to the optimum temperatures for sunflower growth (i.e., Chimenti et al., 2000). However, supra optimum temperature during seed filling could affect weight per seed (Jacum et al., 2000) and oil concentration (Rondanini et al., 2000). The relationships established in this work explained most of variability in weight per seed and oil concentration among different environments and adequately predicted these two yield components in independent experiments. They help to explain the lower weight per seed and oil concentration that are often found in temperate regions when sowing date, a practice with high bal and impact on oil yield (Andrade, 1995; Aguirreza Pereyra, 1998), is delayed (in these regions, incident solar radiation during seed filling of late-sowed crops is often greatly decreased). These relationships are simple

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tools that could be applied to simulate these two oil yield components in sunflower crops. Strictly, these relationships could only be applied to simulate the weight per seed and oil concentration of Dekalb G-100 (black hull and high oil concentration cultivar) grown under good water and mineral conditions with a plant density between 45 000 and 80 000 plants ha1, and good seed set. Oil concentration of a stripped hull and low oil concentration potential cultivar did not vary in response to changes in intercepted radiation during seed filling (Dosio et al., 2000). Nevertheless, several hybrids with black hull and high oil concentration potential have shown a similar response to changes in intercepted radiation to that described by Dosio et al. (2000) for Dekalb G-100 (Izquierdo et al., 1998; Izquierdo, 2000, unpublished data). It could then be expected that application of these relationships to other genotypes with the black hull and high oil concentration would only require recalculation of the equation parameters. Oil yield of the sunflower crop grown under adequate environmental and management conditions could be estimated by coupling these equations with such as seed number estimation by a photothermal quotient (Cantagallo et al., 1997) and leaf area simulation from daily average temperature (i.e., Sadras and Hall, 1988; Dosio et al., 2000).

CONCLUSION
Final weight per seed and oil concentration were largely determined by intercepted radiation from 250 to 450C DAF. Relationships established using these results acceptably predicted the weight per seed and oil concentration data of independent experiments. They helped to explain differences in weight per seed and oil concentration of sunflower crops grown under good moisture and mineral conditions, relatively free of pests and diseases.
ACKNOWLEDGMENTS This work was supported by the Universidad Nacional de Mar del Plata, INTA, Oleaginosa Moreno Hnos. S.A., Nidera S.A., and Aceitera General Deheza S.A. Results presented here were taken from GAAD M.S. Thesis, and M. Ferreiro nomo Thesis. GAAD and YL were and L.M.G Ingeniero Agro fellowships from FOMEC (UNMdP) and INAPG (France), respectively. FHA and LANA are members of the Consejo Naficas y Te cnicas (CONICET, cional de Investigaciones Cient Argentina). Dr. A. Merrien (CETIOM, France) provided useful comments during this work and Dr. O. Turc (INRA MontpellierENSAM, France) made helpful comments on the manuscript. Dr. P. Abbate (INTA) provided the meteorological data. REFERENCES
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