Wide variation of winter-induced sustained thermal energy dissipation in conifers: a common-garden study.

Low temperature in winter depresses rates of photosynthesis, which, in evergreen plants, can exacerbate imbalances between light absorption and photochemical light use. Damage that could result from increased excess light absorption is minimized by the conversion of excitation energy to heat in a process known as energy dissipation, which involves the de-epoxidized carotenoids of the xanthophyll cycle. Overwintering evergreens employ sustained forms of energy dissipation observable even after lengthy periods of dark acclimation. Whereas most studies of photoprotective energy dissipation examine one or a small number of species; here, we measured the levels of sustained thermal energy dissipation of seventy conifer taxa growing outdoors under common-garden conditions at the Red Butte Garden in Salt Lake City, Utah, U.S.A. (forty nine taxa were also sampled for needle pigment content). We observed an extremely wide range of wintertime engagement of sustained energy dissipation; the percentage decrease in dark-acclimated photosystem II quantum efficiency from summer to winter ranged from 6 to 95%. Of the many pigment-based parameters measured, the magnitude of the seasonal decrease in quantum efficiency was most closely associated with the seasonal increase in zeaxanthin content expressed on a total chlorophyll basis, which explained only slightly more than one-third of the variation. We did not find evidence for a consistent wintertime decrease in needle chlorophyll content. Thus, the prevailing mechanism for winter decreases in solar-induced fluorescence emitted by evergreen forests may be decreases in fluorescence quantum yield, and wintertime deployment of sustained energy dissipation likely underlies this effect.

Biochemistry and physiology of foliar isoprene production.

Leaves of some plants emit isoprene, a volatile hydrocarbon. This is formed by a novel chloroplastic isoprenoid biosynthetic pathway, the 1-deoxy-D-xylulose-5-phosphate pathway. The thermoprotection hypothesis suggests that isoprene protects thylakoids from damage at high temperatures. In this article, we discuss the most recent discoveries about the metabolic pathway underlying isoprene biosynthesis, explore the experimental evidence surrounding thermoprotection and advance some alternative hypotheses about the adaptive role that isoprene biosynthesis might play.

Acclimation of Foliar Antioxidant Systems to Growth Irradiance in Three Broad-Leaved Evergreen Species.

The protective role of leaf antioxidant systems in the mechanism of plant acclimation to growth irradiance was studied in Vinca major, Schefflera arboricola, and Mahonia repens, which were grown for several months at 20, 100, and 1200 [mu]mol photons m-2 s-1. As growth irradiance increased, several constituents of the "Mehler-peroxidase" pathway also increased: superoxide dismutase, ascorbate peroxidase, glutathione reductase, ascorbate, and glutathione. This occurred concomitantly with increases in the xanthophyll cycle pool size and in the rate of nonphotochemical energy dissipation under steady-state conditions. There was no evidence for photosystem II overreduction in plants grown at high irradiance, although the reduction state of the stromal NADP pool, estimated from measurements of NADP-malate dehydrogenase activity, was greater than 60% in V. major and S. arboricola. Ascorbate, which removes reactive O2 species generated by O2 photoreduction in the chloroplast and serves as a reductant for the conversion of the xanthophyll cycle pigments to the de-epoxidized forms A plus Z, generally exhibited the most dramatic increases in response to growth irradiance. We conclude from these results that O2 photoreduction occurs at higher rates in leaves acclimated to high irradiance, despite increases in xanthophyll cycle-dependent energy dissipation, and that increases in leaf antioxidants protect against this potential oxidative stress.

Energy dissipation and radical scavenging by the plant phenylpropanoid pathway.

Environmental stresses such as high light, low temperatures, pathogen infection and nutrient deficiency can lead to increased production of free radicals and other oxidative species in plants. A growing body of evidence suggests that plants respond to these biotic and abiotic stress factors by increasing their capacity to scavenge reactive oxygen species. Efforts to understand this acclimatory process have focused on the components of the 'classical' antioxidant system, i.e. superoxide dismutase, ascorbate peroxidase, catalase, monodehydroascorbate reductase, glutathione reductase and the low molecular weight antioxidants ascorbate and glutathione. However, relatively few studies have explored the role of secondary metabolic pathways in plant response to oxidative stress. A case in point is the phenylpropanoid pathway which is responsible for the synthesis of a diverse array of phenolic metabolites such as flavonoids, tannins, hydroxycinnamate esters and the structural polymer lignin. These compounds are often induced by stress and serve specific roles in plant protection, i.e. pathogen defence, ultraviolet screening, antiherbivory, or structural components of the cell wall. This review will highlight a novel antioxidant function for the taxonomically widespread phenylpropanoid metabolite chlorogenic acid (CGA; 5-O-caffeoylquinic acid) and assess its possible role in abiotic stress tolerance. The relationship between CGA biosynthesis and photosynthetic carbon metabolism will also be discussed. Based on the properties of this model phenolic metabolite, we propose that under stress conditions phenylpropanoid biosynthesis may represent an alternative pathway for photochemical energy dissipation that has the added benefit of enhancing the antioxidant capacity of the cell.