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Most environmental stress conditions promote the production of potentially toxic active oxygen species in plant cells. Plants respond with changes in their antioxidant and photoprotective systems. Antioxidants and pigments have been widely used to measure these responses. Because trees are exposed to multiple man-made and natural stresses, their responses are not reflected by changes in single stress markers, but by complex biochemical changes. To evaluate such response patterns, explorative multivariate statistics have been used. In the present study, 12 biochemical variables (chloroplast pigments, state of the xanthophyll cycle, α-tocopherol, ascorbate and dehydroascorbate, glutathione and oxidized glutathione) were measured in previous-year needles of field-grown  Pinus ponderosa  Dougl.  ex  Laws. The trees were sampled in two consecutive years in the San Bernardino Mountains in southern California, where a pollution gradient is overlaid by gradients in natural stresses (drought, altitude). To explore irradiance effects, needle samples were taken directly in the field (sun exposed) and from detached, dark-adapted branches. A principal component analysis on this data set ( n  = 80) resulted in four components (Components 1–4) that explained 67% of the variance in the original data. Component 1 was positively loaded by concentrations of α-tocopherol, total ascorbate and xanthophyll cycle pools, as well as by the proportion of de-epoxides in the xanthophyll cycle. It was negatively loaded by the proportion of dehydroascorbate in the ascorbate pool. Component 2 was negatively loaded by chlorophyll concentrations, and positively loaded by the ratios of lutein and b-carotene to chlorophyll and by the de-epoxidation state of the xanthophyll cycle. Component 3 was negatively loaded by GSH concentrations and positively loaded by the proportions of GSSG and tocopherol concentrations. Component 4 was positively loaded by neoxanthin and negatively loaded by β-carotene. The four components could be assigned to the concerted action of the biochemical protection system: high scores on Component 1 represent highly activated antioxidative defense, changes in pigment composition are represented in Components 2 and 4, and the glutathione system, which is important for antioxidant regeneration, is represented in Component 2. Although Component 1 scores were generally higher (indicating activation of antioxidant defense) in light-adapted needles relative to dark-adapted needles, they were also site dependent with increased scores at sites with less pollution, but higher natural stress impacts. High scores of Components 2 and 3 at the highest elevation site, which was only moderately polluted, indicated an increase in photoprotection by pigments and activation of the glutathione system. Significant differences between light- and dark-adapted needles in Components 2 and 3 were only found at the site with the highest pollution. Use of accumulated variables (components) instead of single biochemical variables enabled recognition of response patterns at particular sites and a better comparison with results of other studies is expected. Typical response patterns could be assigned to particular environmental stress combinations, providing a means of assessing potential biological risks within individual forest stands.

Multivariate patterns of biochemical responses of Pinus ponderosa trees at field plots in the San Bernardino Mountains, southern California