Growth and development at cold-hardening temperatures. Pigment and benzoquinone accumulation in winter rye
Accumulation of chlorophyll, the carotenoids (β-carotene, lutein, violaxanthin, and neoxanthin), and the benzoquinones (plastoquinone A and α-tocopherol) were monitored in 'Puma' rye as a function of leaf ontogeny at warm and cold-hardening temperatures. Although the kinetics of accumulation differed among the leaves of warm-grown plants, the initial and maximum levels of the pigments and benzoquinones expressed on a leaf area basis did not differ significantly among the first four leaves of the main culm. In contrast, the third and fourth leaf of cold-grown plants, which developed completely at the low temperature, generally exhibited initial and maximum pigment and benzoquinone levels 60–300% greater than was observed for leaf 1 and 2 of cold-grown plants, which were completely or partially developed at the warm temperature regime. This resulted in pigment and benzoquinone levels which were 1.6- to 3-fold greater in the plants grown at cold-hardening temperatures than those grown at the warm temperature, when expressed on a per unit leaf area basis. However, when pigment accumulation was calculated on a chlorophyll basis, the benzoquinone content of leaves that developed solely at cold-hardening temperatures exhibited a 1.7-fold increase over the same leaves developed at warm temperatures. Carotenoids did not exhibit this trend. Calculations based on chlorophyll/carotenoid content and dry weight accumulation indicated that leaves that were developed at cold-hardening temperatures appeared to produce more dry matter per unit of photosynthetic pigments than the same leaves that were developed at nonhardening temperatures.