Clouds homogenize shoot temperatures, transpiration, and photosynthesis within crowns of Abies fraseri (Pursh.) Poiret.

Multiple studies have examined the effects of clouds on shoot and canopy-level microclimate and physiological processes; none have yet done so on the scale of individual plant crowns. We compared incident photosynthetically active radiation (PAR), leaf temperatures, chlorophyll fluorescence, and photosynthetic gas exchange of shoots in three different spatial locations of Abies fraseri crowns on sunny (clear to partly cloudy) versus overcast days. The field site was a Fraser fir farm (1038 m elevation) in the Appalachian mountains, USA. Ten saplings of the same age class were marked and revisited for all measurements. Sunny conditions corresponded with 5-10× greater sunlight incidence on south-facing outer shoots compared to south-facing inner and north-facing outer shoots, which were shaded and received only indirect (diffuse) sunlight. Differences in spatial distribution of irradiance were mirrored in differences in shoot temperatures, photosynthesis, and transpiration, which were all greater in south-facing outer shoots compared to more shaded crown locations. In contrast, overcast conditions corresponded with more homogeneous sunlight distribution between north and south-facing outer shoots, and similar shoot temperatures, chlorophyll fluorescence (ΦPSII), photosynthesis, and transpiration; these effects were observed in south-facing inner shoots as well, but to a lesser extent. There was no significant difference in conductance between different crown locations on sunny or overcast days, indicating spatial differences in transpiration under sunny conditions were likely driven by leaf temperature differences. We conclude that clouds can affect spatial distribution of sunlight and associated physiological parameters not only within forest communities, but within individual crowns as well.

Photosynthetic costs and benefits of abaxial versus adaxial anthocyanins in Colocasia esculenta 'Mojito'.

MAIN CONCLUSION: Anthocyanins in upper (adaxial) leaf tissues provide greater photoprotection than in lower (abaxial) tissues, but also predispose tissues to increased shade acclimation and, consequently, reduced photosynthetic capacity. Abaxial anthocyanins may be a compromise between these costs/benefits. Plants adapted to shaded understory environments often exhibit red/purple anthocyanin pigmentation in lower (abaxial) leaf surfaces, but rarely in upper (adaxial) surfaces. The functional significance of this color pattern in leaves is poorly understood. Here, we test the hypothesis that abaxial anthocyanins protect leaves of understory plants from photo-oxidative stress via light attenuation during periodic exposure to high incident sunlight in the forest understory, without interfering with sunlight capture and photosynthesis during shade conditions. We utilize a cultivar of Colocasia esculenta exhibiting adaxial and abaxial anthocyanin variegation within individual leaves to compare tissues with the following color patterns: green adaxial, green abaxial (GG), green adaxial, red abaxial (GR), red adaxial, green abaxial (RG), and red adaxial, red abaxial (RR). Consistent with a photoprotective function of anthocyanins, tissues exhibited symptoms of increasing photoinhibition in the order (from least to greatest): RR, RG, GR, GG. Anthocyanic tissues also showed symptoms of shade acclimation (higher total chl, lower chl a/b) in the same relative order. Inconsistent with our hypothesis, we did not observe any differences in photosynthetic CO2 uptake under shade conditions between the tissue types. However, GG and GR had significantly (39 %) higher photosynthesis at saturating irradiance (A sat) than RG and RR. Because tissue types did not differ in nitrogen content, these patterns likely reflect differences in resource allocation at the tissue level, with greater nitrogen allocated toward energy processing in GG and GR, and energy capture in RG and RR (consistent with relative sun/shade acclimation). We conclude that abaxial anthocyanins are likely advantageous in understory environments because they provide some photoprotection during high-light exposure, but without the cost of decreased A sat associated with adaxial anthocyanin-induced shade syndrome.

Cloud immersion: an important water source for spruce and fir saplings in the southern Appalachian Mountains.

Cloud immersion can provide a potentially important moisture subsidy to plants in areas of frequent fog including the threatened spruce-fir communities of the southern Appalachian Mountains (USA). These mountaintop communities grow only above ~1,500 m elevation, harbor the endemic Abies fraseri, and have been proposed to exist because of frequent cloud immersion. While several studies have demonstrated the importance of cloud immersion to plant water balance, no study has evaluated the proportion of plant water derived from cloud moisture in this ecosystem. Using the isotopic mixing model, IsoSource, we analyzed the isotopic composition of hydrogen and oxygen for water extracted from ground water, deep soil, shallow soil, fog, and plant xylem at the upper and lower elevational limits both in May (beginning of the growing season) and October (end of the growing season). Cloud-immersion water contributed up to 31% of plant water at the upper elevation sites in May. High-elevation plants of both species also experienced greater cloud immersion and had greater cloud water absorption (14-31%) compared to low-elevation plants (4-17%). Greater cloud water uptake occurred in May compared to October, despite similar rainfall and cloud-immersion frequencies. These results demonstrate the important water subsidy that cloud-immersion water can provide. With a warming climate leading potentially to increases in the ceiling of the cloud base and, thus, less frequent cloud immersion, persistence of these relic mountaintop forests may depend on the magnitude of these changes and the compensating capabilities of other water sources.

Support for a photoprotective function of winter leaf reddening in nitrogen-deficient individuals of Lonicera japonica.

Plants growing in high-light environments during winter often exhibit leaf reddening due to synthesis of anthocyanin pigments, which are thought to alleviate photooxidative stress associated with low-temperature photoinhibition through light attenuation and/or antioxidant activity. Seasonal high-light stress can be further exacerbated by a limited photosynthetic capacity, such as nitrogen-deficiency. In the present study, we test the following hypotheses using three populations of the semi-evergreen vine Lonicera japonica: (1) nitrogen deficiency corresponds with reduced photosynthetic capacity; (2) individuals with reduced photosynthetic capacity synthesize anthocyanin pigments in leaves during winter; and (3) anthocyanin pigments help alleviate high-light stress by attenuating green light. All populations featured co-occurring winter-green and winter-red leafed individuals on fully-exposed (high-light), south-facing slopes in the Piedmont of North Carolina, USA. Consistent with our hypotheses, red leaves consistently exhibited significantly lower foliar nitrogen than green leaves, as well as lower total chlorophyll, quantum yield efficiency, carboxylation efficiency, and photosynthesis at saturating irradiance (Asat). Light-response curves measured using ambient sunlight versus red-blue LED (i.e., lacking green wavelengths) demonstrated significantly reduced quantum yield efficiency and a higher light compensation point under sunlight relative to red-blue LED in red leaves, but not in green leaves, consistent with a (green) light-attenuating function of anthocyanin pigments. These results are consistent with the hypothesis that intraspecific anthocyanin synthesis corresponds with nitrogen deficiency and reduced photosynthetic capacity within populations, and support a light-attenuating function of anthocyanin pigments.

Lessons from the re-emergence of imported measles cases following the COVID-19 pandemic in Victoria, Australia.

Introduction: Australia was declared to have eliminated endemic measles in 2014; however, imported cases continue to pose a threat of outbreaks. International travel restrictions during the coronavirus disease 2019 (COVID-19) pandemic led to a rapid decline in measles cases. The re-opening of the Australian international border to measles endemic regions returns the threat of outbreaks, which may be further compounded by disruptions in routine vaccinations during the COVID-19 pandemic. We consider lessons learned from the public health response to recent measles cases. Methods: This case series includes all confirmed measles cases meeting the national case definition reported to the Victorian Government Department of Health (the Department) between 1 January and 31 December 2022. The Department conducted active case finding and contact tracing of all cases in line with national guidelines. Cases were descriptively analysed. Results: In 2022, six of the seven measles cases reported in Australia occurred in Victoria, all of whom resided in Australia and acquired their infection overseas. Three cases were unlinked, and three formed an epidemiologically-linked household cluster. One case was partially vaccinated, one was not eligible for vaccination, one had unknown vaccination status, and three were unvaccinated, one of whom was under 12 months old but would have been eligible for vaccination prior to travel to endemic regions. None of the cases led to secondary transmission within Australia. Discussion: Following the COVID-19 pandemic, measles importations have re-commenced in Victoria. Although few measles cases occurred in 2022 and none resulted in onwards transmission, imported measles cases remain complex and require substantial public health follow-up. Delays in case diagnosis and flight contact tracing pose a significant risk for outbreaks of measles. Public health interventions are needed to maintain high vaccination rates, improve contact tracing, and ensure public health authorities and healthcare providers can rapidly identify and respond to imported measles cases.

Red leaf margins indicate increased polygodial content and function as visual signals to reduce herbivory in Pseudowintera colorata.

Red-pigmented leaf margins are common, but their functional significance is unknown. We hypothesized that red leaf margins reduce leaf herbivory by signalling to herbivorous insects the presence of increased chemical defences. Leaves were collected from a natural population of Pseudowintera colorata. Margin size, herbivory damage, anthocyanin content and concentrations of polygodial, a sesquiterpene dialdehyde with antifeedant properties, were quantified. Feeding trials involving larvae of Ctenopseustis obliquana, a generalist herbivore, were conducted on red- and green-margined P. colorata leaves in darkness, or under white, green or red light. Leaves with wider red margins contained higher concentrations of polygodial and anthocyanins, and incurred less natural herbivory. In trials under white light, C. obliquana consumed disproportionately more green- than red-margined leaf laminae. Larvae exhibited no feeding preference when light was manipulated such that leaf colour discrimination was impaired. Red leaf margins provide a reliable and effective visual signal of chemical defence in P. colorata. Ctenopseustis obliquana larvae perceive and respond to the colour of the leaf margins, rather than to olfactory signals. Our study provides direct experimental evidence for aposematic coloration in red leaves.

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.

Winter leaf reddening in 'evergreen' species.

Leaf reddening during autumn in senescing, deciduous tree species has received widespread attention from the public and in the scientific literature, whereas leaf reddening in evergreen species during winter remains largely ignored. Winter reddening can be observed in evergreen herbs, shrubs, vines and trees in Mediterranean, temperate, alpine, and arctic regions, and can persist for several months before dissipating with springtime warming. Yet, little is known about the functional significance of this colour change, or why it occurs in some species but not others. Here, the biochemistry, physiology and ecology associated with winter leaf reddening are reviewed, with special focus on its possible adaptive function. Photoprotection is currently the favoured hypothesis for winter reddening, but alternative explanations have scarcely been explored. Intraspecific reddening generally increases with sunlight incidence, and may also accompany photosynthetic inferiority in photosynthetically 'weak' (e.g. low-nitrogen) individuals. Red leaves tend to show symptoms of shade acclimation relative to green, consistent with a photoprotective function. However, winter-red and winter-green species often cohabitate the same high-light environments, and exhibit similar photosynthetic capacities. The factors dictating interspecific winter leaf colouration therefore remain unclear. Additional outstanding questions and future directions are also highlighted, and possible alternative functions of winter reddening discussed.

The same anthocyanins served four different ways: Insights into anthocyanin structure-function relationships from the wintergreen orchid, Tipularia discolor.

Over 500 unique anthocyanins have been described to date, which vary in color, antioxidant, light-attenuating, and antimicrobial properties. Identification of anthocyanin chemical structure may therefore serve as an important clue to their in situ function in plants. We characterized the histological and chemical structures of anthocyanins associated with diverse leaf color patterns in the terrestrial orchid, Tipularia discolor, as a step towards understanding their ultimate function. Tipularia discolor produces a single wintergreen leaf in autumn, which is drab brown in color during expansion. Upper (adaxial) surfaces of fully-expanded leaves may be green, purple-spotted, or solid purple, while lower (abaxial) surfaces are bright magenta. Our results showed that the same three cyanidin 3,7,3'-triglucosides, in similar concentrations and proportions, accounted for coloration in each of these cases, and that different colors result from differences in histological location of anthocyanins (i.e. abaxial/adaxial epidermis, mesophyll). Anthocyanins with 3,7,3' linkage positions are rare in plants, occurring only within the orchid subfamily Epidendroideae, to which Tipularia belongs. These results are important to the discussion of anthocyanin structure-function because they serve as a reminder that 1) plants may employ the same anthocyanins in different anatomical locations to achieve a broad range of colors (and potentially adaptive functions), and 2) anthocyanin chemical structure and anatomical location are influenced by phylogenetic inertia, as well as natural selection.

Association between winter anthocyanin production and drought stress in angiosperm evergreen species.

Leaves of many evergreen angiosperm species turn red under high light during winter due to the production of anthocyanin pigments, while leaves of other species remain green. There is currently no explanation for why some evergreen species exhibit winter reddening while others do not. Conditions associated with low leaf water potentials (Psi) have been shown to induce reddening in many plant species. Because evergreen species differ in susceptibility to water stress during winter, it is hypothesized that species which undergo winter colour change correspond with those that experience/tolerate the most severe daily declines in leaf Psi during winter. Six angiosperm evergreen species which synthesize anthocyanin in leaves under high light during winter and five species which do not were studied. Field Psi, pressure/volume curves, and gas exchange measurements were derived in summer (before leaf colour change had occurred) and winter. Consistent with the hypothesis, red-leafed species as a group had significantly lower midday Psi in winter than green-leafed species, but not during the summer when all the leaves were green. However, some red-leafed species showed midday declines similar to those of green-leafed species, suggesting that low Psi alone may not induce reddening. Pressure-volume curves also provided some evidence of acclimation to more negative water potentials by red-leafed species during winter (e.g. greater osmotic adjustment and cell wall hardening on average). However, much overlap in these physiological parameters was observed as well between red and green-leafed species, and some of the least drought-acclimated species were red-leafed. No difference was observed in transpiration (E) during winter between red and green-leaved species. When data were combined, only three of the six red-leafed species examined appeared physiologically acclimated to prolonged drought stress, compared to one of the five green-leafed species. This suggests that drought stress alone is not sufficient to explain winter reddening in evergreen angiosperms.

Red (anthocyanic) leaf margins do not correspond to increased phenolic content in New Zealand Veronica spp.

BACKGROUND AND AIMS: Red or purple coloration of leaf margins is common in angiosperms, and is found in approx. 25 % of New Zealand Veronica species. However, the functional significance of margin coloration is unknown. We hypothesized that anthocyanins in leaf margins correspond with increased phenolic content in leaf margins and/or the leaf entire, signalling low palatability or leaf quality to edge-feeding insects. METHODS: Five species of Veronica with red leaf margins, and six species without, were examined in a common garden. Phenolic content in leaf margins and interior lamina regions of juvenile and fully expanded leaves was quantified using the Folin-Ciocalteu assay. Proportions of leaf margins eaten and average lengths of continuous bites were used as a proxy for palatability. KEY RESULTS: Phenolic content was consistently higher in leaf margins compared with leaf interiors in all species; however, neither leaf margins nor more interior tissues differed significantly in phenolic content with respects to margin colour. Mean phenolic content was inversely correlated with the mean length of continuous bites, suggesting effective deterrence of grazing. However, there was no difference in herbivore consumption of red and green margins, and the plant species with the longest continuous grazing patterns were both red-margined. CONCLUSIONS: Red margin coloration was not an accurate indicator of total phenolic content in leaf margins or interior lamina tissue in New Zealand Veronica. Red coloration was also ineffective in deterring herbivory on the leaf margin, though studies controlling for variations in leaf structure and biochemistry (e.g. intra-specific studies) are needed before more precise conclusions can be drawn. It is also recommended that future studies focus on the relationship between anthocyanin and specific defence compounds (rather than general phenolic pools), and evaluate possible alternative functions of red margins in leaves (e.g. antioxidants, osmotic adjustment).

Optical effects of abaxial anthocyanin on absorption of red wavelengths by understorey species: revisiting the back-scatter hypothesis.

A red/purple coloration of lower (abaxial) leaf surfaces is commonly observed in deeply-shaded understorey plants, especially in the tropics. However, the functional significance of red abaxial coloration, including its role in photosynthetic adaptation, remains unclear. The objective of this study was to test the back-scatter hypothesis for abaxial leaf coloration, which posits that red pigments internally reflect/scatter red light transmitted by the upper leaf surface back into the mesophyll, thereby enhancing photon capture in light-limited environments. Abaxially red/non-red variegated leaves of Begonia heracleifolia (Cham. & Schltdl.) were used to compare reflectance spectra and chlorophyll fluorescence profiles of abaxially anthocyanic (red) and acyanic (non-red) tissues under red light. Photosynthetic gas exchange in response to red light was also compared for abaxially red/non-red leaf sections. The results did not support a back-scattering function, as anthocyanic leaf surfaces were not more reflective of red light than acyanic surfaces. Anthocyanic tissues also did not exhibit any increases in the mesophyll absorbance of red light, or increased photosynthetic gas exchange under red light at any intensity, relative to acyanic tissues. These results suggest that abaxial anthocyanins do not significantly enhance the absorption of red light in the species tested, and alternative functions are discussed.

Student performance on the Test of Scientific Literacy Skills (TOSLS) does not change with assignment of a low-stakes grade.

OBJECTIVE: Response-validated multiple-choice assessments are used in college courses to assess student learning gains. The ability of a test to accurately reflect student learning gains is highly dependent on the students' effort. Within our institution, lackluster student effort is common on response-validated multiple-choice concept assessments that are not included as a portion of the semester grade but are used to inform curricular changes. Thus, we set out to determine whether increasing testing stakes by assigning a grade on student performance had an effect on student score and self-reported effort. The Test of Scientific Literacy Skills (TOSLS) is a response-validated multiple-choice assessment used to measure scientific literacy in undergraduates. We administered the TOSLS to students enrolled in a general education Biology course, both during the first 2 weeks (pretest) and the last 2 weeks (posttest) of the course. RESULTS: Self-reported effort and TOSLS performance were significantly correlated in the ungraded cohort. This relationship did not exist in the graded sections. Our data indicate that assigning a low-stakes grade has no significant effect on mean student performance or self-reported effort on the TOSLS within our general education course.

Importance of natural cloud regimes to ecophysiology in the alpine species, Caltha leptosepala and Arnica parryi, Snowy Range Mountains, southeast Wyoming, USA.

The south-central Rocky Mountains, USA, are characterised by a dry, continental mesoclimate with typical convective cloud formation during the afternoon. Little is known about the specific influence of such predictable cloud patterns on the microclimate and ecophysiology of associated species. During the summer of 2012, days with afternoon clouds were most common (50% of all days) compared with completely clear (24%) or cloudy days (6.5%). In two representative alpine species, Caltha leptosepala DC. and Arnica parryi A. Gray, fully overcast days reduced mean daily photosynthesis (A) by nearly 50% relative to fully clear days. Mean afternoon A was significantly lower on fully cloudy days relative to days with afternoon clouds only or no clouds in both species. Notably, A did not differ during afternoon cloud days relative to clear afternoons. Afternoon clouds significantly reduced transpiration (E) in C. leptosepala relative to clear days, and both species showed mean reductions in plant water stress (i.e. higher Ψ), though this difference was not significant. Water use efficiency (WUE) (A/E) decreased from morning to afternoon, especially on cloudy days, and the presence of clouds had a positive effect on the light reactions of photosynthesis based on fluorescence measurements (Fv'/Fm'), in both species. Cloudy days were characterised by higher Fv/Fm than afternoon clouds and clear days during both the morning and the afternoon (especially for A. parryi) and recovery to near pre-dawn values for cloudy and afternoon cloud day types, but not clear days. Overall, similar ecophysiological advantages of this typical afternoon cloud pattern was apparent in both species, although their spatial microsite differences related to winter snow accumulation may also play an important role.

Estimating contribution of anthocyanin pigments to osmotic adjustment during winter leaf reddening.

The association between plant water stress and synthesis of red, anthocyanin pigments in leaves has led some plant biologists to propose an osmotic function of leaf reddening. According to this hypothesis, anthocyanins function as a solute in osmotic adjustment (OA), contributing to depression of osmotic potential (Ψ(π)) and maintenance of turgor pressure during drought-stressed conditions. Here we calculate the percent contribution of anthocyanin to leaf Ψ(π) during OA in two angiosperm evergreen species, Galax urceolata and Gaultheria procumbens. Both species exhibit dramatic leaf reddening under high light during winter, concomitant with declines in leaf water potential and accumulation of solutes. Data previously published by the authors on osmotic potential at full turgor (Ψ(π,100)) of G. urceolata and G. procumbens leaves before and after leaf reddening were used to estimate OA. In vivo molar concentrations of anthocyanin, glucose, fructose, and sucrose measured from the same individuals were converted to pressure equivalents using the Ideal Gas Law, and percent contribution to OA was estimated. Estimated mean OA during winter was -0.7MPa for G. urceolata and -0.8MPa for G. procumbens. In vivo concentrations of anthocyanin (3-10mM) were estimated to account for ∼2% of OA during winter, and comprised <0.7% of Ψ(π,100) in both species. Glucose, fructose, and sucrose combined accounted for roughly 50 and 80% of OA for G. urceolata and G. procumbens, respectively, and comprised ∼20% of Ψ(π,100). We observed that a co-occurring, acyanic species (Vinca minor) achieved similar OA without synthesizing anthocyanin. We conclude that anthocyanins represent a measurable, albeit meager, component of OA in red-leafed evergreen species during winter. However, due to their low concentrations, metabolic costliness relative to other osmolytes, and striking red color (unnecessary for an osmotic function), it is unlikely that they are synthesized solely for an osmoprotectant role.

Unravelling the evolution of autumn colours: an interdisciplinary approach.

Leaf colour change is commonly observed in temperate deciduous forests in autumn. This is not simply a side effect of leaf senescence, and, in the past decade, several hypotheses have emerged to explain the evolution of autumn colours. Yet a lack of crosstalk between plant physiologists and evolutionary ecologists has resulted in slow progress, and so the adaptive value of this colour change remains a mystery. Here we provide an interdisciplinary summary of the current body of knowledge on autumn colours, and discuss unresolved issues and future avenues of research that might help reveal the evolutionary meaning of this spectacle of nature.

Seasonal photosynthesis and anthocyanin production in 10 broadleaf evergreen species.

Leaves of many evergreen species turn red when exposed to high sunlight during winter due to production of photoprotective anthocyanin pigments, while leaves of other species, lacking anthocyanin, remain green. Why some evergreen species synthesise anthocyanin pigments while others do not is currently unknown. Furthermore, the relative photosynthetic performance of anthocyanic (red) and acyanic (green) evergreens has yet to be described. Here we present seasonal ecophysiological data for five red and green broadleaf evergreen species. We hypothesise that species which synthesise anthocyanins in winter leaves correspond to those with the most drastic seasonal photosynthetic declines, as reduced energy sinks increase vulnerability to photoinhibition and need for photoprotection. Our results did not support this hypothesis, as gas exchange measurements showed no difference in mean seasonal photosynthetic capacity between red- and green-leafed species. Consistent with anthocyanin's shading effect, red-leafed species had significantly higher chlorophyll content, lower chlorophyll a/b ratios, and higher maximum light capture efficiency of PSII (Fv/Fm) than green-leafed species during the winter, but not during the summer (when all leaves were green). We conclude that anthocyanin production during winter is likely not associated with diminished photosynthetic capacity, and may simply represent an alternative photoprotective strategy utilised by some species during winter.

Attenuation of incident light in Galax urceolata (Diapensiaceae): concerted influence of adaxial and abaxial anthocyanic layers on photoprotection.

Although anthocyanin coloration in lower (abaxial) leaf cells has been documented for numerous species, the functional significance of this character has not been comprehensively investigated according to habitat or leaf orientation. Here, we demonstrate that abaxial anthocyanin may function as a photoprotectant, similarly to its purported role in upper (adaxial) cells, in leaves vulnerable to high irradiance incident on abaxial surfaces. Spectral scans were derived for Galax urceolata leaves with the following phenotypes: abaxial or adaxial anthocyanin only, abaxial and adaxial anthocyanin, and no anthocyanin. To determine whether anthocyanins conferred protection from photoinhibition, maximum photosystem II efficiencies of red (anthocyanic) and green (acyanic) surfaces were compared during and after exposure to photoinhibitory conditions. Leaves were either positioned with their adaxial surfaces facing the light source or inverted to expose abaxial surfaces. Spectral scans showed increased absorptance of 500-600 nm wavelengths by red surfaces (consistent with the absorbance spectrum of anthocyanin), regardless of whether that surface was abaxial or adaxial. Leaves with anthocyanin in either illuminated surface were also photoinhibited less than leaves lacking anthocyanin in that surface. These results suggest that anthocyanic layers reduce absorbed sunlight in the mesophyll not only for adaxial surfaces, but also for the abaxial. Adaxial/abaxial anthocyanin plasticity may therefore be adaptive in high-light environments or during light-sensitive developmental stages where leaf orientation and/or substrate albedo are variable.

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species.

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.