Growth patterns and biomass spectra

The growth cycle of small-grain cereals involves changes in size, form and number of plant organs. The external stages of cereal growth include germination, crop emergence, seedling growth, tillering, stem elongation, booting, inflorescence emergence, anthesis and maturity (Fig. 1). The classical monitoring of crop biomass requires destructive samplings of plants at different growth stages, counting of the number of plants contained in the sample and its weighing after oven-drying them. Crop biomass may be expressed as crop dry weight (CDW), which can be obtained from the plants sampled at a given stage as the product of average dry weight per plant (W, g) and the number of plants per unit area, and is frequently expressed as g m-2 (Villegas et al., 2001). The leaf area expansion of a cereal crop may be monitored through changes in its leaf area index (LAI, a dimensionless value), which is the ratio of leaf green area to the area of ground on which the crop is growing. LAI may be calculated as the product of the mean one-sided leaf area per plant (LAP, m2 plant-1) and the number of plants per unit area in the sample (plants m-2). Changes in total green area of the crop may be described through the green area index (GAI, a dimensionless value), which is the ratio of total green area of the plants (leaves and stems, as well as spike peduncles and spikes when applicable) to the area of ground on which the crop is growing. It can be calculated as the product of total green area per plant (GAP, m2 plant-1) and the number of plants per unit area in the sample (plants m-2) (Royo et al., 2004).

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Fig. 1. Growth stages of small-grain cereals. Numbers correspond to the Zadoks scale (Zadoks et al., 1974)

Raw data from destructive sampling can be fitted to mathematical models, usually empirically based, to describe the growth pattern during the crop cycle. The logistic model of Richards (Richards, 1959), the expolinear equation of Goudriaan & Monteith (Goudriaan & Monteith, 1990), and the asymmetric logistic peak curve first used by Royo and Tribo (Royo & Tribo, 1997), have been used to describe the growth of crops. This last model has been useful for monitoring the biomass and leaf area expansion of triticale (Royo & Blanco,

1999) and durum wheat (Royo et al., 2004; Villegas et al., 2001). The mathematical models present the variation in dry matter production, leaf area or green area expansion over time, allowing variations between species (Fig. 2), genotypes, years and environmental conditions to be assessed (Fig. 3). Similarly to the case of grain yield, variability induced by the genetic background in the growth pattern of small-grain cereals has been found to be lower than the environmental variation caused by either year or site effects (Royo et al., 2004; Villegas et al.,

2001) .

Crop growth conditions can be monitored by measuring the spectra reflected by crop canopies in the visible (VIS, A=400-700 nm) and near-infrared (NIR, A =700-1300 nm) regions of the electromagnetic spectrum (Fig. 4). Given that the amount of green area of a canopy determines the absorption of photosynthetic active radiation by photosynthetic organs, spectral reflectance measurements can provide an instantaneous quantitative assessment of
the crop’s ability to intercept radiation and photosynthesize (Ma et al., 1996). Therefore, the absorption by the crop canopy of very specific wavelengths of electromagnetic radiation is associated with certain morphological and physiological crop attributes related to the development of the total photosynthetic area of the canopy.

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Подпись: Fig. 2. Illustration of the differences between the patterns of biomass accumulation and leaf area expansion of barley (A), spring triticale (□), and winter triticale (•) from experiments conducted in 4 Mediterranean environments. Samples were taken at seedling (S), tillering (T), beginning of jointing (J), booting (B), anthesis (A), and physiological maturity (M). Biomass increased continually from anthesis to maturity in barley, but in triticale the peak of biomass took place between anthesis and maturity. The maximum LAI was reached at the booting stage in barley, but a little later in triticale. Adapted from Royo & Tribo (1997)

Days from sowing

The reflectance spectra of a healthy crop-canopy shows a relative maximum around 550 nm, a relative minimum around 680 nm and an abrupt increase around 700 nm, remaining fairly constant beyond this point (Fig. 4). The spectral reflectance in the VIS wavelengths depends on the absorption of incident radiation by leaf chlorophyll and associated pigments such as

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carotenoid and anthocyanins. Crop reflectance is very low in the blue (400-500 nm) and red (600-700 nm) regions of the spectrum, because they contain the peaks of chlorophyll absorbance. Beyond 700 nm the reflectance of the NIR wavelengths is high since it is not absorbed by plant pigments and is scattered by plant tissues at different levels in the canopy (Knipling, 1970).

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Wavelength (nm)

Visible Near-infrared

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Blue Green Red

Fig. 4. Variation of the reflectance spectra of a healthy wheat canopy at different growth stages compared with the bare soil spectrum. H, heading; A, anthesis; M, milk-grain stage; PM, physiological maturity. The magnitude of the increase in reflectance at around 700 nm indicates differences in biomass