Photosynthesis and Photoprotection in Overwintering Plants

Abstract: Seasonal differences in the capacity of photosynthetic electron transport, leaf pigment composition, xanthophyll cycle characteristics and chlorophyll fluorescence emission were investigated in two biennial mesophytes ( Malva neglecta and Verbascum thapsus ) that grow in full sunlight, and in leaves/needles of sun and shade populations of several broad-leafed evergreens and conifers (Vinca minor, Euonymus kiautschovicus, Mahonia repens, Pseudotsuga menziesii [Douglas fir], and Pinus ponderosa). Both mesophytic species maintained or upregulated photosynthetic capacity in the winter and exhibited no upregulation of photoprotection. In contrast, photosynthetic capacity was downregulated in sun leaves/needles of V. minor, Douglas fir, and Ponderosa pine, and even in shade needles of Douglas fir. Interestingly, photosynthetic capacity was upregulated during the winter in shade leaves/needles of V. minor, Ponderosa pine and Euonymus kiautschovicus. Nocturnal retention of zeaxanthin and antheraxanthin, and their sustained engagement in a state primed for energy dissipation, were observed largely in the leaves/needles of sun-exposed evergreen species during winter. Factors that may contribute to these differing responses to winter stress, including chloroplast redox state, the relative levels of source and sink activity at the whole plant level, and apoplastic versus symplastic phloem loading, are discussed.     

The leaf anatomy of a broad-leaved evergreen allows an increase in leaf nitrogen content in winter

In temperate regions, evergreen species are exposed to large seasonal changes in air temperature and irradiance. They change photosynthetic characteristics of leaves responding to such environmental changes. Recent studies have suggested that photosynthetic acclimation is strongly constrained by leaf anatomy such as leaf thickness, mesophyll and chloroplast surface facing the intercellular space, and the chloroplast volume. We studied how these parameters of leaf anatomy are related with photosynthetic seasonal acclimation. We evaluated differential effects of winter and summer irradiance on leaf anatomy and photosynthesis. Using a broad-leaved evergreen Aucuba japonica , we performed a transfer experiment in which irradiance regimes were changed at the beginning of autumn and of spring. We found that a vacant space on mesophyll surface in summer enabled chloroplast volume to increase in winter. The leaf nitrogen and Rubisco content were higher in winter than in summer. They were correlated significantly with chloroplast volume and with chloroplast surface area facing the intercellular space. Thus, summer leaves were thicker than needed to accommodate mesophyll surface chloroplasts at this time of year but this allowed for increases in mesophyll surface chloroplasts in the winter. It appears that summer leaf anatomical characteristics help facilitate photosynthetic acclimation to winter conditions. Photosynthetic capacity and photosynthetic nitrogen use efficiency were lower in winter than in summer but it appears that these reductions were partially compensated by higher Rubisco contents and mesophyll surface chloroplast area in winter foliage.     

Contrasting patterns of photosynthetic acclimation and photoinhibition in two evergreen herbs from a winter deciduous forest

The relationship between the microclimate within an Oak-Hickory forest and photosynthetic characters of two resident evergreen herbs with contrasting leaf phenologies was investigated on a monthly basis for 1 full year. Heuchera americana has leaf flushes in the spring and fall, with average leaf life spans of 6–7 months. Hexastylis arifolia produces a single cohort of leaves each spring with a leaf life span of 12–13 months. We predicted that among evergreen plants inhabiting a seasonal habitat, a species for which the frequency of leaf turnover is greater than the frequency of seasonal extremes would have a greater annual range in photosynthetic capacity than a species that only produced a single flush of leaves during the year. Photosynthetic parameters, including apparent quantum yield, maximum photosynthetic capacity (P_max), temperature of maximum photosynthesis, photochemical efficiency of PSII and leaf nitrogen (N) and chlorophyll concentrations, were periodically measured under laboratory conditions in leaves sampled from natural populations of both species. Mature leaves of both species acclimated to changing understory conditions with the mean seasonal differences being significantly greater for Heuchera than for Hexastylis. Area based maximum photosynthetic rates at 25°C were approximately 250% and 100% greater in winter leaves than summer leaves for Heuchera and Hexastylis respectively. Nitrogen concentrations were highest in winter leaves. Chlorophyll concentrations were highest in summer leaves. Low P_max/N values for these species suggest preferential allocation of leaf nitrogen into non-photosynthetic pools and/or light-harvesting function at the expense of photosynthetic enzymes and electron transport components. Despite the increase in photosynthetic capacity, there was evidence of chronic winter photoinhibition in Hexastylis, but not in Heuchera. Among these ecologically similar species, there appears to be a trade-off between the frequency of leaf production and the balance of photosynthetic acclimation and photoinhibition.     

Annual photosynthetic activities of temperate evergreen and deciduous broadleaf tree species with simultaneous and successive leaf emergence in response to altitudinal air temperature

To explain why the composition of evergreen and deciduous forests changes along air temperature gradients, we measured several traits of single leaves from temperate deciduous and evergreen broadleaf trees with simultaneous and successive leaf emergence growing at different altitudes in the field. The parameters included seasonal net photosynthetic rate, longevity, mass per area, nitrogen content, and photosynthetic nitrogen-use efficiency. With decreasing altitude, the leaf longevity of deciduous broadleaf trees increased, whereas the maximum net photosynthetic rate decreased. In contrast, leaf longevity of evergreen broadleaf trees decreased, whereas the minimum net photosynthetic rate in winter increased. Along the air temperature gradient, the annual production of deciduous trees with simultaneous leaf emergence may be constant, because the integrated lifetime net photosynthetic rate (ILNPR) of a single leaf changed little. In comparison, deciduous trees with successive leaf emergence may show enhanced annual production with increasing air temperature, by increasing the total leaf number per branch and tree under an extended growing season. Temperate evergreen broadleaf tree species may also show increased annual production with increasing air temperature by sufficiently raising the number of the first-year leaves to the total leaves of branch and tree, which is accelerated by raising the integrated first-year net photosynthetic rate of the single leaf, despite little change in the ILNPR. With increasing air temperature from cool-temperate to warm-temperate zones, evergreen broadleaf tree species gain an advantage of the annual production over deciduous broadleaf tree species with simultaneous leaf emergence.     

哀牢山和猫儿山中山常绿和落叶阔叶树光合特性对季节温度变化的响应

研究调查了云南哀牢山中山常绿阔叶林和广西猫儿山中山落叶水青冈林中共生的常绿和落叶阔叶树光合特性在2005年的季节变化,探讨了两地温度气候差异对植物优势度的影响。在夏季,同一生境下的常绿植物和落叶植物之间以及两个生境树种之间的光合速率没有显著差别。当温度下降时,两个生境内的常绿和落叶植物的光合速率都下降,这与气孔的部分关闭和光系统Ⅱ活性的下降有显著的关系。气孔的部分关闭有利于减少水分蒸腾损失,但落叶树气孔关闭程度明显高于常绿植物而且无法避免严重的光抑制。常绿植物通过气孔调节来提高光合水分利用效率和通过光系统Ⅱ活性调节来降低光合机构的损伤来维持叶片四季常绿,但是猫儿山中山森林冬季低温严重损害气孔和光系统Ⅱ的功能而使常绿植物生长受到严重制约并限制其在群落中成为优势种,与之相反,哀牢山中山森林冬季相对暖和导致常绿植物能积累更多的光合产物并促进其在群落中形成优势。     

Autumn photosynthetic decline and growth cessation in seedlings of white spruce are decoupled under warming and photoperiod manipulations

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high‐latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down‐regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change.     

Do gymnosperm needles pull water through the xylem produced in the same year as the needle?

This research investigated the longevity of functional connections between leaf traces and stem xylem in 16 species of conifers to better understand the spatial use of sapwood for water transport. The first question was which ring(s) stained when a vacuum was applied to the distal end of the cut surface of a needle attached to a short stem segment. The vacuum was applied to either 1- or 2-yr-old foliage taken from 4-6-yr-old saplings. The 16 species were then categorized based on the growth ring that most consistently stained when the vacuum was applied to the 2-yr-old needles. There were three distinct stain patterns for the evergreen conifers and one pattern for the deciduous conifers. Three evergreen species could not be categorized. The second question used needle fall data to ask whether the leaf trace appeared to break because of age or stem diameter in 5-yr-old Pseudotsuga menziesii saplings. An apparent threshold stem diameter at which needles tended to be shed was more related to diameter than age. These xylem connection patterns could affect leaf cohort physiology and the spatial pattern of water flux in sapwood.     

Intermittent low temperatures constrain spring recovery of photosynthesis in boreal Scots pine forests

During winter and early spring, evergreen boreal conifers are severely stressed because light energy cannot be used when photosynthesis is pre‐empted by low ambient temperatures. To study photosynthetic performance dynamics in a severe boreal climate, seasonal changes in photosynthetic pigments, chloroplast proteins and photochemical efficiency were studied in a Scots pine forest near Zotino, Central Siberia. In winter, downregulation of photosynthesis involved loss of chlorophylls, a twofold increase in xanthophyll cycle pigments and sustained high levels of the light stress‐induced zeaxanthin pigment. The highest levels of xanthophylls and zeaxanthin did not occur during the coldest winter period, but rather in April when light was increasing, indicating an increased capacity for thermal dissipation of excitation energy at that time. Concomitantly, in early spring the D1 protein of the photosystem II (PSII) reaction centre and the light‐harvesting complex of PSII dropped to their lowest annual levels. In April and May, recovery of PSII activity, chloroplast protein synthesis and rearrangements of pigments were observed as air temperatures increased above 0°C. Nevertheless, severe intermittent low‐temperature episodes during this period not only halted but actually reversed the physiological recovery. During these spring low‐temperature episodes, protective processes involved a complementary function of the PsbS and early light‐induced protein thylakoid proteins. Full recovery of photosynthesis did not occur until the end of May. Our results show that even after winter cold hardening, photosynthetic activity in evergreens responds opportunistically to environmental change throughout the cold season. Therefore, climate change effects potentially improve the sink capacity of boreal forests for atmospheric carbon. However, earlier photosynthesis in spring in response to warmer temperatures is strongly constrained by environmental variation, counteracting the positive effects of an early recovery process.     

Coordination of leaf N,anatomy, photosynthetic capacity,and hydraulics enhances evergreen expansive potential

Key message

Reduced leaf longevity, N-fixation, and enhanced hydraulic capacity combined support greater shifts in seasonal photosynthetic capacity of an expansive understory evergreen woody species relative to co-occurring less expansive evergreen species.

Abstract

Physiological functioning typically declines with increased leaf life span. While an evergreen leaf habit is generally associated with reduced leaf N, physiological capacity, and slower growth, most expansive woody species are evergreens and/or N fixers. An evergreen leaf habit enables year-round activity and less investment in carbon and nutrients, while N-fixation enhances photosynthetic capacity. Our objective was to compare anatomy and physiology of three woody evergreens Ilex opaca Aiton (Aquifoliaceae), Kalmia latifolia L. (Ericaceae), and Myrica cerifera (Myricaceae) of varying leaf longevity, N-fixation capability, and known expansive potential in a deciduous forest understory to determine if seasonal physiological performance integrated these factors. We hypothesized that I. opaca (non-expansive) and K. latifolia (moderately expansive), which have longer leaf longevities, would have reduced physiological performance compared to M. cerifera (expansive), which has shorter leaf longevity, and symbiotically fixes atmospheric N. Stomatal conductance to water vapor, photosynthetic and hydraulic capacities, specific leaf area, and leaf %N decreased with increasing leaf life span; however, trends among species were not consistent seasonally. While hydraulic capacity remained constant throughout the year, photosynthetic capacity did not. During the growing season, M. cerifera displayed photosynthetic capacity similar to deciduous species, yet, during the winter, photosynthetic capacity was similar to the slower-growing evergreens. Reduced leaf life span, enhanced hydraulic capacity, and nitrogen fixation support the seasonal shift in photosynthetic capacity observed in M. cerifera. This “hybrid” strategy enables M. cerifera to maximize productivity during months of optimal conditions, thereby promoting rapid growth and expansion in the understory.     

Photosynthesis of Conifers in Relation to Annual Growth Cycles and Dry Matter Production

Observations that deciduous larch species can show annual growth increments equal to or greater than evergreen conifers, and that the saturating light intensity for photosynthesis in needles of Larix leptolepis was almost twice those for several evergreen conifers, led to a study of the photosynthetic mechanism in L. leptolepis. Several features of photosynthesis in L. leptolepis placed this species in an intermediate position between classical C₃ and C₄ plants. Incorporation of ¹⁴C from ¹⁴CO₂ by enzyme preparations of larch needles was eight times greater with PEP as substrate than with ribulose bis phosphate; a chlorophyll a/b ratio of 3.5 was obtained; needles possessed a green starch-containing endodermis but with little orientation of mesophyll cells to this “bundle sheath”; no clear ultrastructural dimorphism was observed between chloroplasts of mesophyll and endodermal cells; a CO₂-compensation point of 20 μl-l^?1 was recorded; and the first measurable product of photosynthesis appeared to be malate rather than phospho-glyceric acid. These results are discussed in relation to the deciduous habit of L. leptolepis and its high productivity in comparison with other conifers.     

Evolutionary significance of a flat-leaved Pinus in Vietnamese rainforest

Pines are generally absent from tropical rainforests. An important exception, Pinus krempfii, is a unique tree that bears flattened needles and competes with evergreen angiosperm trees in southern Vietnam. Here, the photosynthetic and hydraulic physiology of P. krempfii leaves were examined to determine whether this species departs from the widespread pattern of high-light-demanding photosynthetic physiology displayed in needle-leaved Pinus species. Maximum photosynthesis and light saturation of photosynthesis, as well as stem and leaf hydraulic efficiencies, were all very low in P. krempfii compared with other Pinus species. These characteristics were consistent with our observations of P. krempfii seedling regeneration under the forest canopy. By possessing shade tolerance coupled with the production of flattened leaves, P. krempfii has converged morphologically and physiologically with many genera of the southern hemisphere conifer family Podocarpaceae. This convergence extends to a key feature of leaf anatomy, the production of tubular sclereids in the leaf for radial transport of water from the vein to the margin. These observations suggest that few adaptive possibilities are open to conifers when moving into tropical rainforest, meaning that Pinus is forced into direct competition with southern hemisphere conifers for a narrow niche in the equatorial zone.     

Foliar carbon isotope discrimination in Larix species and sympatric evergreen conifers: a global comparison

Larches (Larix spp.), deciduous conifers, occur in the northern hemisphere in cold-temperate and boreal climates – an environment normally thought to favor evergreen tree species. We compare foliar carbon isotope discrimination (Δ), instantaneous water use efficiency, total foliar nitrogen concentration, and specific leaf area (for a subset of sites) between Larix spp. and co-occurring evergreen conifers at 20 sites throughout the natural range of larches. Except for Larix occidentalis in the xeric Intermountain West, USA, Δ is significantly (P < 0.05) greater for larches than co-occurring evergreen conifers at 77% of the sites, suggesting that larches use water less efficiently. At elevations greater than 3000 m, the Δ of Larix spp. and co-occurring conifers converge, suggesting that water is not the limiting resource. Foliar nitrogen concentration and specific leaf area are two ecophysiological characteristics that are positively correlated with high photosynthetic capacity. Foliar nitrogen concentration is significantly greater for larches than evergreen conifers at 88% of the sites and specific leaf area is approximately three times greater for larches than co-occurring conifers. Future studies should examine the potential effect that global warming may have on the distribution of larch forests because the water use efficiency of larches is commonly less than co-occurring evergreen conifers and the boreal and high-latitude environments are likely to experience the greatest climate warming. Received: 23 May 1997 / Accepted: 28 October 1997     

Freezing Tolerance,Pigments and SOD of Five Conifers in Shenyang

Cultivated Picea koraiensis Nakai, Pinus sylvestris var. mongolica Litvin, P. koraiensis Sieb et Zuce, P. tabulaeformis Carr. and P. bungeana Zucc. were studied. Changes in chlorophylls carotenoids contents, SOD activities and iso-zymograms were monitored for the needles of the 5 evergreen conifers and related to their freezing tolerance during winter of 1.987–1988 in Shenyang. It shows that the chlorophylls were degradated to various extent in the winter while the carotenoids were still synthesized during the soil-frozen period. SOD activities increased from July to November for all 5 conifers and the more tolerate the species, the higher the SOD activities during winter. The results suggest that SOD possibly has important contribution to freezing tolerance besides protecting the needles from winter photooxidation. SOD iso-zymograms and their variation showed differance among species as well, but it is hard to draw a generalized conclussion.     

Estimating photosynthetic rate and annual carbon gain in conifers from specific leaf weight and leaf biomass

Summary Canopy photosynthesis is difficult to measure directly or to predict with complex models demanding knowledge of seasonal variation in environmental and physiological properties of the canopy. Trees in particular offer a challenge with their large, aerodynamically rough and seasonally-changing canopy properties. In this paper we assess the possibility of using specific leaf weight to predict seasonal and annual net photosynthetic rate in deciduous (Larix sp.) and evergreen (Picea abies) conifers.Annual photosynthetic rate and specific leaf weight of different positions of the crown in both species were highly correlated (r ²=0.930). Annual carbon uptake by different segments in a mature P. abies crown was closely related to leaf biomass. The relationship was improved by adjusting the leaf biomass of each segment in regard to its specific leaf weight relative to the maximum found in the canopy. The adjustment accounted for associated differences in photosynthetic activity. This combined structural index (leaf biomassxrelative specific leaf weight) could, when calibrated, predict the total annual carbon uptake by different parts of the crown. If direct measurements of photosynthesis are not available, the combined structural index may still serve as a comparative estimator of annual carbon uptake.     

Leaf phenology and carbon dynamics in six leguminous trees

I documented photosynthetic rates and seasonal stem total nonstructural carbohydrates (TNC) in six leguminous tree species Burkea africana, Baikiaea plurijuga, Erythrophleum africanum, Guibourtia coleosperma, Julbernardia globiflora and Pterocarpus angolensis exhibiting a range of leaf phenological patterns. My goal was to (i) measure photosynthetic characteristics and levels of stored stem carbohydrates in species with varying patterns of leaf phenology and (ii) determine seasonal patterns of stem carbohydrate storage. Despite significant differences in the timing of bud break and leaf cover between the six species, there were no significant differences in maximum photosynthetic rate, quantum efficiency or light saturation point between species. Similarly, there was no significant difference in seasonal mean stem TNC levels despite significant differences in the timing of bud break and leaf cover both between species and within a single species. However, while the average amount of TNC does not seem to be related to leaf phenology, the patterns of carbohydrate use and storage do seem to be related to leaf phenology.     

Effects of artificial frost hardening and winter stress on net photosynthesis, photosynthetic electron transport and RuBP carboxylase activity in seedlings of Pinus silvestris

Net photosynthesis of seedlings of Pinus silvestris has been measured and compared with the activities of photosynthetic electron transport and extracted RuBP carboxylase. The effects of prolonged frost hardening (photoperiod 8 h, + 3°C) followed by winter stress at subzero temperatures were analysed. There was a parallel effect of frost hardening and winter stress on the photosynthetic properties of both intact seedlings and isolated chloroplast thylakoids. The activity of extracted RuBP carboxylase was less affected by the treatments. In relation to earlier works we conclude that the decay of net photosynthesis in winter climate is determined by the electron transport properties of the chloroplast thylakoids, i.e. by the pool sizes of photosynthetically active plastoquinone. The results of this work justify the definition of two phases in the response of conifers towards autumn and winter climates: I. Frost hardening occurs at temperatures slightly above zero and it does not affect the efficiency of photosynthesis as defined by the quantum yield at rate limiting light absorption. II. Winter stress occurs at subzero temperatures and it is characterized by a suppression of the photosynthetic efficiency as a result of damage within the photosynthetic apparatus.     

Maximum CO2-assimilation rates of vascular plants on an Alaskan arctic tundra slope

Light-saturated CO₂-assimilation rates of 19 vascular plant species were measured on a tundra slope in the foothills of the Brooks Range, Alaska. Maximum assimilation capacities on a leaf area basis ranged from 20.3 μmol m⁻² s⁻¹ for the forb, Bistorta plumosa , to 6.0 μmol m⁻² s⁻¹ for the evergreen, Empetrum hermaphroditicum . Graminoids, deciduous shrubs, and forbs fell within a similar range of maximum photosynthetic rates on a leaf area basis. Evergreens had the lowest rates. On a leaf weight basis, maximum assimilation rates were greatest for forbs, followed by deciduous shrubs, graminoids, and evergreens. Rates of evergreens were less than half those of all other growth forms. Cassiope tetragona had the lowest rates per unit leaf weight of any species tested; mean maximum rates of C. tetragona were only 14% of those of B. plumosa , the species with the highest rates. When the data were subjected to canonical analysis, only a partial correspondence was found between species growth form and photosynthetic characteristics.     

Enhanced ethylene emissions from red and norway spruce exposed to acidic mists

Acidic cloudwater is believed to cause needle injury and to decrease winter hardiness in conifers. During simulations of these adverse conditions, rates of ethylene emissions from and levels of 1-aminocyclopropane-1-carboxylic acid (ACC) in both red and Norway spruce needles increased as a result of treatment with acidic mists but amounts of 1-malonyl(amino)cyclopropane-1-carboxylic acid remained unchanged. However, release of significant quantities of ethylene by another mechanism independent of ACC was also detected from brown needles. Application of exogenous plant growth regulators such as auxin, kinetin, abscisic acid and gibberellic acid (each 0.1 millimolar) had no obvious effects on the rates of basal or stress ethylene production from Norway spruce needles. The kinetics of ethylene formation by acidic mist-stressed needles suggest that there is no active inhibitive mechanism in spruce to prevent stress ethylene being released once ACC has been formed.     

Diurnal and seasonal variations in photosynthetic and morphological traits of the tree ferns Dicksonia antarctica (Dicksoniaceae) and Cyathea australis (Cyatheaceae) in wet sclerophyll forests of Australia

Steady state and dynamic responses of two tree fern species of contrasting origins, Dicksonia antarctica (of Gondwanan origin) and Cyathea australis (Pan-tropical), were studied over two consecutive years under field conditions in a wet sclerophyll forest of south-east Australia. Irrespective of their different origins, there were no significant differences in photosynthetic performance between the two species. Growth irradiance and leaf temperature, but not plant water status, was significantly related to photosynthetic and morphological traits. At a common leaf temperature, maximum light-use efficiency of photosystem II (F_v/F_m) was significantly lower in winter than in summer, suggesting some limitation to PSII efficiency potentially associated with cold winter mornings. Both species displayed seasonal acclimation in a number of measured photosynthetic parameters and frond traits (i.e. F_v/F_m, A_sat, g_s, N_A, total chlorophyll, SLA). Acclimation of stomatal density to spatial variation in growth irradiance seemed limited in both species, although stomatal pattern differed between species. Because there were no significant differences between the two species in photosynthetic parameters, both species can be described by common carbon gain and water use models at the leaf scale.     

Xanthophyll cycle pigment and antioxidant profiles of winter-red (anthocyanic) and winter-green (acyanic) angiosperm evergreen species

Leaves of many angiosperm evergreen species change colour from green to red during winter, corresponding with the synthesis of anthocyanin pigments. The ecophysiological function of winter colour change (if any), and why it occurs in some species and not others, are not yet understood. It was hypothesized that anthocyanins play a compensatory photoprotective role in species with limited capacity for energy dissipation. Seasonal xanthophyll pigment content, chlorophyll fluorescence, leaf nitrogen, and low molecular weight antioxidants (LMWA) of five winter-red and five winter-green angiosperm evergreen species were compared. Our results showed no difference in seasonal xanthophyll pigment content (V+A+Z g(-1) leaf dry mass) or LMWA between winter-red and winter-green species, indicating red-leafed species are not deficient in their capacity for non-photochemical energy dissipation via these mechanisms. Winter-red and winter-green species also did not differ in percentage leaf nitrogen, corroborating previous studies showing no difference in seasonal photosynthesis under saturating irradiance. Consistent with a photoprotective function of anthocyanin, winter-red species had significantly lower xanthophyll content per unit chlorophyll and less sustained photoinhibition than winter-green species (i.e. higher pre-dawn F(v)/F(m) and a lower proportion of de-epoxidized xanthophylls retained overnight). Red-leafed species also maintained a higher maximum quantum yield efficiency of PSII at midday (F'(v)/F'(m)) during winter, and showed characteristics of shade acclimation (positive correlation between anthocyanin and chlorophyll content, and negative correlation with chlorophyll a/b). These results suggest that the capacity for photon energy dissipation (photochemical and non-photochemical) is not limited in red-leafed species, and that anthocyanins more likely function as an alternative photoprotective strategy to increased VAZ/Chl during winter.