Leaf Xanthophyll content and composition in sun and shade determined by HPLC

As a part of our investigations to test the hypothesis that zeaxanthin formed by reversible de-epoxidation of violaxanthin serves to dissipate any excessive and potentially harmful excitation energy we determined the influence of light climate on the size of the xanthophyll cycle pool (violaxanthin + antheraxanthin + zeaxanthin) in leaves of a number of species of higher plants. The maximum amount of zeaxanthin that can be formed by de-epoxidation of violaxanthin and antheraxanthin is determined by the pool size of the xanthophyll cycle. To quantitate the individual leaf carotenoids a rapid, sensitive and accurate HPLC method was developed using a non-endcapped Zorbax ODS column, giving baseline separation of lutein and zeaxanthin as well as of other carotenoids and Chl a and b.The size of the xanthophyll cycle pool, both on a basis of light-intercepting leaf area and of light-harvesting chlorophyll, was ca. four times greater in sun-grown leaves of a group of ten sun tolerant species than in shade-grown leaves in a group of nine shade tolerant species. In contrast there were no marked or consistent differences between the two groups in the content of the other major leaf xanthophylls, lutein and neoxanthin. Also, in each of four species examined the xanthophyll pool size increased with an increase in the amount of light available during leaf development whereas there was little change in the content of the other xanthophylls. However, the -carotene/-carotene ratio decreased and little or no -carotene was detected in sun-grown leaves. Among shade-grown leaves the -carotene/-carotene ratio was considerably higher in species deemed to be umbrophilic than in species deemed to be heliophilic.The percentage of the xanthophyll cycle pool present as violaxanthin (di-epoxy-zeaxanthin) at solar noon was 96–100% for shade-grown plants and 4–53% for sun-grown plants with zeaxanthin accounting for most of the balance. The percentage of zeaxanthin in leaves exposed to midday solar radiation was higher in those with low than in those with high photosynthetic capacity.The results are consistent with the hypothesis that the xanthophyll cycle is involved in the regulation of energy dissipation in the pigment bed, thereby preventing a buildup of excessive excitation energy at the reaction centers.Abbreviations A antheraxanthin - C -carotene - C -carotene - EPS epoxidation state (V+0.5A)/(V+A+Z) - L lutein - N neoxanthin - PFD photon flux density - V violaxanthin - Z zeaxanthin C.I.W.-D.P.B. Publiation No. 1035     

Inclination of sun and shade leaves influences chloroplast light harvesting and utilization

Light harvesting and utilization by chloroplasts located near the adaxial vs the abaxial surface of sun and shade leaves were examined by fluorometry in two herbaceous perennials that differed in their anatomy and leaf inclination. Leaves of Thermopsis montana had well-developed palisade and spongy mesophyll whereas the photosynthetic tissue of Smilacina stellata consisted of spongy mesophyll only. Leaf orientation depended upon the irradiance during leaf development. When grown under low-light levels, leaves of S. stellata and T. montana were nearly horizontal, whereas under high-light levels, S. stellata leaves and T. montana leaves were inclined 60⁰ and 30⁰, respectively. Leaf inclination increased the amount of light that was intercepted by the lower leaf surfaces and affected the photosynthetic properties of the chloroplasts located near the abaxial leaf surface. The slowest rates of quinone pool reduction and reoxidation were found in chloroplasts located near the adaxial leaf surface of T. montana plants grown under high light, indicating large quinone pools in these chloroplasts. Chloroplasts near the abaxial surface of low-light leaves had lower light utilization capacities as shown by photochemical quenching measurements. The amount of photosystem II (PSII) down regulation, measured from each leaf surface, was also found to be influenced by irradiance and leaf inclination. The greatest difference between down regulation monitored from the adaxial vs abaxial surfaces was found in plants with horizontal leaves. Different energy dissipation mechanisms may be employed by the two species. Values for down regulation in S. stellata were 2–3 times higher than those in T. montana, while the portion of the PSII population which was found to be Q_B nonreducing was 4–6 times lower in high light S. stellata leaves than in T. montana. All values of Stern-Volmer type nonphotochemical quenching (NPQ) from S. stellata leaves were similar when quenching analysis was performed at actinic irradiances that were higher than the irradiance to which the leaf surface was exposed during growth. In contrast, with T. montana, NPQ values from the abaxial leaf surface were up to 45% higher than those from the adaxial leaf surface regardless of growth conditions. The observed differences in chloroplast properties between species and between the adaxial and abaxial leaf surfaces may depend upon a complex interaction among light, leaf anatomy and leaf inclination.     

Responses of photosystem I compared with photosystem II to high-light stress in tropical shade and sun leaves

Sun and shade leaves of several plant species from a neotropical forest were exposed to excessive light to evaluate the responses of photosystem I in comparison to those of photosystem II. Potential photosystem I activity was determined by means of the maximum P700 absorbance change around 810 nm (ΔA_810max) in saturating far-red light. Leaf absorbance changes in dependence of increasing far-red light fluence rates were used to calculate a ‘saturation constant’, K_s, representing the far-red irradiance at which half of the maximal absorbance change (ΔA_810max/2) was reached in the steady state. Photosystem II efficiency was assessed by measuring the ratio of variable to maximum chlorophyll fluorescence, F_v/F_m, in dark-adapted leaf samples. Strong illumination caused a high degree of photo-inhibition of photosystem II in all leaves, particularly in shade leaves. Exposure to 1800–2000 μ mol photons m⁻² s⁻¹ for 75 min did not substantially affect the potential activity of photosystem I in all species tested, but caused a more than 40-fold increase of K_s in shade leaves, and a three-fold increase of K_s in sun leaves. The increase in K_s was reversible during recovery under low light, and the recovery process was much faster in sun than in shade leaves. The novel effect of high-light stress on the light saturation of P700 oxidation described here may represent a complex reversible mechanism within photosystem I that regulates light-energy dissipation and thus protects photosystem I from photo-oxidative damage. Moreover, we show that under high-light stress a high proportion of P700 accumulates in the oxidized state, P700⁺. Presumably, conversion of excitation energy to heat by this cation radical may efficiently contribute to photoprotection.     

Xanthophyll cycle components and capacity for non-radiative energy dissipation in sun and shade leaves ofLigustrum ovalifolium exposed to conditions limiting photosynthesis

The relationships between photosynthetic efficiency, non-radiative energy dissipation and carotenoid composition were studied in leaves ofLigustrum ovalifolium developed either under full sunlight or in the shade. Sun leaves contained a much greater pool of xanthophyll cycle components than shade leaves. The rate of non-radiative energy dissipation, measured as non-photochemical fluorescence quenching (NPQ), was strictly related to the deepoxidation state (DPS) of xanthophyll cycle components in both sun and shade leaves, indicating that zeaxanthin (Z) and antheraxanthin (A) are involved in the development of NPQ. Under extreme conditions of excessive energy, sun leaves showed higher maximum DPS than shade leaves. Therefore, sun leaves contained not only a greater pool of xanthophyll cycle components but also a higher proportion of violaxanthin (V) actually photoconvertible to A and Z, compared to shade leaves. Both these effects contributed to the higher NPQ in sun versus shade leaves. The amount of photoconvertible V was strongly related to chla/b ratio and inversely to leaf neoxanthin content. This evidence indicates that the amount of photoconvertible V may be dependent on the degree of thylakoid membrane appression and on the organization of chlorophyll-protein complexes, and possible explanations are discussed. Exposure to chilling temperatures caused a strong decline in the photon yield of photosynthesis and in the intrinsic efficiency of PS II photochemistry in sun leaves, but little effects in shade leaves. These effects were accompanied by increases in the pool of xanthophyll cycle components and in DPS, more pronounced in sun than in shade leaves. This corroborates the view that Z and A may play a photoprotective role under unfavorable conditions. In addition to the xanthophyll-related non-radiative energy dissipation, a slow relaxing component of NPQ, independent from A and Z concentrations, has been found in leaves exposed to low temperature and high light. This quenching component may be attributed either to other regulatory mechanism of PS II efficiency or to photoinactivation.Research supported by National Research Council of Italy, Special Project RAISA, Sub-Project 2, Paper N. 1587.     

不同年龄麻竹阴阳叶生态生理特性

麻竹是中国重要的大型经济竹种,其栽培已从过去河滩、四旁零散种植发展到规模化培育,通过山地麻竹发笋期内不同年龄植株阴阳叶养分和代谢动态的比较研究,结果表明麻竹阳叶氮素、磷素浓度比阴叶高,但钾素浓度阳叶低于阴叶;从发笋初期至末期阴阳叶氮、磷、钾素浓度都呈逐渐减少的变化趋势,阴阳叶氮、磷、钾素浓度差异逐渐减小;阳叶在净光合速率、暗呼吸速率、CO2补偿点、光补偿点、光饱和点等方面较阴叶高,光呼吸较低,但不同年龄麻竹之间各指标变化有所不同.     

Developmental process of sun and shade leaves in Chenopodium album L.

The authors’ previous study of Chenopodium album L. revealed that the light signal for anatomical differentiation of sun and shade leaves is sensed by mature leaves, not by developing leaves. They suggested that the two‐cell‐layered palisade tissue of the sun leaves would be formed without a change in the total palisade tissue cell number. To verify that suggestion, a detailed study was made of the developmental processes of the sun and shade leaves of C. album with respect to the division of palisade tissue cells (PCs) and the data was expressed against developmental time (leaf plastochron index, LPI). The total number of PCs per leaf did not differ between the sun and shade leaves throughout leaf development (from LPI ?1 to 10). In both sun and shade leaves, anticlinal cell division of PCs occurred most frequently from LPI ?1 to 2. In sun leaves, periclinal division of PCs occurred synchronously with anticlinal division. The constancy of the total number of PCs indicates that periclinal divisions occur at the expense of anticlinal divisions. These results support the above suggestion that two‐cell‐layered palisade tissue is formed by a change of cell division direction without a change in the total number of PCs. PCs would be able to recognize the polarity or axis that is perpendicular to the leaf plane and thereby change the direction of their cell divisions in response to the light signal from mature leaves.     

Chlorophyll and light gradients in sun and shade leaves of Spinacia oleracea

Abstract. Light gradients were measured and correlated with chlorophyll concentration and anatomy of leaves in spinach (Spinacia oleracea L.). Light gradients were measured at 450, 550 and 680 nm within thin (455 μm) and thick (630 μm) leaves of spinach grown under sun and shade conditions. The light gradients were relatively steep in both types of leaves and 90% of the light at 450 and 680 nm was absorbed by the initial 140 μm of the palisade. In general, blue light was depleted faster than red light which, in turn was depleted faster than green light. Light penetrated further into the thicker palisade of sun leaves in comparison to the shade leaves. The distance that blue light at 450 nm travelled before it became 90% depleted was 120 μm in sun leaves versus 76 μm in shade leaves. Red light at 680 nm and green light at 550 nm travelled further but the trends were similar to that measured at 450nm. The steeper light gradients within the palisade-of shade leaves were caused by increased scattering of light within the intercellular air spaces and/or cells which were less compact than those in sun leaves. The decline in the amount of light within the leaf appeared to be balanced by a gradient in chlorophyll concentration measured in paradermal sections. Progressing from the adaxial epidermis, chlorophyll content increased through the palisade and then declined through the spongy mesophyll. Chlorophyll content was similar in the palisade of both sun and shade leaves. Chloroplast distribution within both sun and shade leaves was relatively uniform so that the chlorophyll gradient appeared to be caused by greater amounts of chlorophyll within chloroplasts located deeper within the leaf. These results indicate that the anatomy of the palisade may be of special importance for controlling the penetration of photo-synthetically active radiation into the leaf. Changing the structural characteristics of individual palisade cells or their arrangement may be an adaptation that maximizes the absorption of light in leaves with varying mesophyll thickness due to different ambient light regimes.     

Carotenoid composition of peridermal twigs does not fully conform to a shade acclimation hypothesis

The photosynthetic pigments of twigs in five tree and shrub species possessing chlorenchyma under a well developed, stomata-less, and highly photon absorptive periderm were analysed and compared to those of the corresponding canopy leaves. We asked whether the unavoidable shade acclimation of corticular chlorenchyma results in photosynthetic pigment complements typically found in shade leaves. As expected, chlorophyll (Chl) a/b ratios in twigs were consistently low. However, carotenoid (Car) analysis did not confirm the initial hypothesis, since twigs generally contained increased Chl-based pool sizes of the xanthophyll cycle components. The contents of photo-selective neoxanthin and lutein were high as well. Yet, -carotene content was extraordinarily low. In addition, twigs retained high pre-dawn ratios of the deepoxidized antheraxanthin and zeaxanthin, although environmental conditions were not pre-disposing for such a state. The unexpected Car composition allows the conclusion that other micro-environmental conditions within twigs (hypoxia, increased red to blue photon ratios, and extremely high CO₂ concentrations) are more important than shade in shaping the Car profiles.This revised version was published online in March 2005 with corrections to the page numbers.     

Relationships between carotenoid composition and growth habit in British plant species

The pigment composition of leaves from a number of different plant species collected from field sites in the region of Sheffield, UK, have been compared using high-performance liquid chromatography. Expression of pigment content per unit leaf area was dominated by variation in the total leaf chlorophyll. Neither chlorophyll per unit area nor the chlorophyll a/b ratio were found to be correlated with the habitat from which the plants originated. When the amounts of different carotenoids were expressed relative to the total carotenoid pool, it was found that whilst neither total carotene (α- +β-carotene) nor neoxanthin correlated with ability to grow in shade, the leaf content of both lutein and the total xanthophyll cycle carotenoids (zeaxanthin, anther-axanthin and violaxanthin) did, with lutein content being high in shade species and xanthophyll cycle intermediates low. There was a strong negative correlation between the relative amounts of each of these groups of carotenoids. The ratio of lutein to xanthophyll cycle carotenoids was strongly correlated to an index of shade tolerance.     

High carbon dioxide and sun/shade effects on isoprene emission from oak and aspen tree leaves

Abstract. Isoprene (2-methyl 1, 3-butadiene) is emitted from many plants, especially trees. We tested the effect of growth at high CO₂ partial pressure and sun versus shade conditions on the capacity of Quercus rubra L. (red oak) and Populus tremuloides Michx. (quaking aspen) leaves to make isoprene. Oak leaves grown at high CO₂ partial pressure (65 Pa) had twice the rate of isoprene emission as leaves grown at 40Pa CO₂. However, aspen leaves behaved oppositely, with high CO₂-grown leaves having just 60-70% the rate of isoprene emission as leaves grown in 40 Pa CO₂. Similar responses were observed from 25 to 35 °C leaf temperature during assay. The stimulation of isoprene emission by growth at high CO₂ and the stimulation in high temperature resulted in isoprene emission consuming over 15% of the carbon fixed during photosynthesis in high-CO₂ grown oak leaves assayed at 35 °C. Leaves from the south (sunny) sides of trees growing in natural conditions had rates of isoprene emission double those of leaves growing in shaded locations on the same trees. This effect was similar in both aspen and oak. The leaves used for these experiments had significantly different chlorophyll a/b ratios indicating they were functionally sun (from the sunny locations) or shade leaves (from the protected locations). Because the metabolic pathway of isoprene synthesis is unknown, we are unable to speculate about how or why these effects occur. However, these effects are more consistent with metabolic control of isoprene release rather than a metabolic leak of isoprene from metabolism. The results are also important for large scale modelling of isoprene emission and for predicting the effect of future increases in atmospheric CO₂ level on isoprene emission from vegetation.     

The spatial distribution of chloroplast water in Acer platanoides sun and shade leaves

We evaluated a new, two-dimensional (2-D) nuclear magnetic resonance (NMR) imaging technique as a method for measuring the distribution of chloroplasts in leaves. NMR images that showed the distribution of chloroplast water and of total water as a function of depth into Acer platanoides sun and shade leaves were compared with the distribution of chlorophyll in the same leaf types (as measured by fluorescence microscopy), with the cellular structure (by scanning electron microscopy), and with published information. Results showed that the volume fraction of chloroplast water was much larger in shade than in sun leaves, and that it averaged about one-third larger in the palisade than in the spongy parenchyma region of both leaf types. Chlorophyll fluorescence was more intense in shade than in sun leaves. In sun leaves, fluorescence was maximal in the palisade region near the junction with the spongy parenchyma, while in shade leaves, fluorescence was maximal in the upper part of the spongy layer. We concluded that 2-D NMR imaging reliably indicates the location of chloroplast water.     

Morpho‐anatomical and physiological differences between sun and shade leaves in Abies alba Mill. (Pinaceae,Coniferales): a combined approach

Morphology, anatomy and physiology of sun and shade leaves of Abies alba were investigated and major differences were identified, such as sun leaves being larger, containing a hypodermis and palisade parenchyma as well as possessing more stomata, while shade leaves exhibit a distinct leaf dimorphism. The large size of sun leaves and their arrangement crowded on the upper side of a plagiotropic shoot leads to self‐shading which is explainable as protection from high solar radiation and to reduce the transpiration via the lamina. Sun leaves furthermore contain a higher xanthophyll cycle pigment amount and Non‐Photochemical Quenching (NPQ) capacity, a lower amount of chlorophyll b and a total lower chlorophyll amount per leaf, as well as an increased electron transport rate and an increased photosynthesis light saturation intensity. However, sun leaves switch on their NPQ capacity at rather low light intensities, as exemplified by several parameters newly measured for conifers. Our holistic approach extends previous findings about sun and shade leaves in conifers and demonstrates that both leaf types of A. alba show structural and physiological remarkable similarities to their respective counterparts in angiosperms, but also possess unique characteristics allowing them to cope efficiently with their environmental constraints.     

植物叶黄素循环的组成、功能和调节(综述)

对近几年来植物体内叶黄素循环的组成、功能、堇菜黄素脱环氧化酶和玉米黄素环氧化酶的结构、生化性质和调节,以及叶黄素的可转变性、定位等方面的研究进展作了综述。     

Carotenoid distribution and deepoxidation in thylakoid pigment-protein complexes from cotton leaves and bundle-sheath cells of maize

In response to excess light, the xanthophyll violaxanthin (V) is deepoxidized to zeaxanthin (Z) via antheraxanthin (A) and the degree of this deepoxidation is strongly correlated with dissipation of excess energy and photoprotection in PS II. However, little is known about the site of V deepoxidation and the localization of Z within the thylakoid membranes. To gain insight into this problem, thylakoids were isolated from cotton leaves and bundle-sheath strands of maize, the pigment protein-complexes separated on Deriphat gels, electroeluted, and the pigments analyzed by HPLC. In cotton thylakoids, 30% of the xanthophyll cycle pigments were associated with the PS I holocomplex, including the PS I light-harvesting complexes and PS I core complex proteins (CC I), and about 50% with the PS II light-harvesting complexes (LHC II). The Chl was evenly distributed between PS I and PS II. Less than 2% of the neoxanthin, about 18% of the lutein, and as much as 76% of the -carotene of the thylakoids were associated with PS I. Exposure of pre-darkened cotton leaves to a high photon flux density for 20 min prior to thylakoid isolation caused about one-half of the V to be converted to Z. The distribution of Z among the pigment-protein complexes was found to be similar to that of V. The distribution of the other carotenoids was unaffected by the light treatment. Similarly, in field-grown maize leaves and in the bundle-sheath strands isolated from them, about 40% of the V present at dawn had been converted to Z at solar noon. Light treatment of isolated bundle-sheath strands which initially contained little Z caused a similar degree of conversion of V to Z. As in cotton thylakoids, about 30% the V+A+Z pool in bundle-sheath thylakoids from maize was associated with the PS I holocomplex and the CC I bands and 46% with the LHC II bands, regardless of the extent of deepoxidation. These results demonstrate that Z is present in PS I as well as in PS II and that deepoxidation evidently takes place within the pigment-protein complexes of both photosystems.Abbreviations A antheraxanthin - CC I, CC II Core or reaction center complex of PS I, PS II - CP Chl protein - EPS epoxidation state - F_m Chl fluorescence at closed PS II reaction centers - IEF isoelectric focussing gels - LHC I, LHC II light-harvesting complex of PS I, PS II - OE oxygen evolving polypeptide - PFD photon flux density - PS I^* PS I holocomplex - V violaxanthin - Z zeaxanthin - antibody against C.I.W.-D.P.B. Publication No. 1127.     

Response of mitochondria to light intensity in the leaves of sun and shade species

The present authors have shown previously that both respiration rates and in vivo activities of the alternative oxidase (AOX) of leaves of Alocasia odora, a shade species, are lower than those in sun species, thereby optimizing energy production under limited light conditions (Noguchi et al., Australian Journal of Plant Physiology 28, 27–35, 2001). In the present study, mitochondria isolated from A. odora leaves were examined in order to investigate the biochemical basis for the differences in respiratory parameters. Alocasia odora and spinach plants were cultivated under both high and low light intensities, mitochondria were isolated from their leaves, and their respiratory properties compared. Mitochondrial content of leaf extracts from the two species was estimated using fumarase activities and antibody detection of porin (the voltage-dependent anion channel of the outer mitochondrial membrane). On a mitochondrial protein basis, spinach leaves showed higher capacities of the cytochrome pathway and cytochrome c oxidase (COX) than A. odora leaves. However, on a mitochondrial protein basis, A. odora showed higher capacities of AOX, which had a high affinity for ubiquinone when activated by pyruvate. Alocasia odora also had larger amounts of mitochondrial protein per leaf dry weight, even under severely shaded conditions, than spinach. Lower growth light intensity led to lower activities of most pathways and proteins tested in both species, especially glycine-dependent oxygen uptake. In the low light environment, most of the AOX protein in A. odora leaves was in its inactive, oxidized dimer form, but was converted to its reduced active form when plants were grown under high light. This shift may prevent over-reduction of the respiratory chain under photo-oxidative conditions.     

Carotenoids and Environmental Stress in Plants: Significance of Carotenoid-Mediated Modulation of Membrane Physical Properties

Carotenoids, apart of their antenna function in photosynthesis, play an important role in the mechanisms protecting the photosynthetic apparatus against various harmful environmental factors. They protect plants against overexcitation in strong light and dissipate the excess of absorbed energy, they scavenge reactive oxygen species formed during photooxidative stress and moderate the effect of extreme temperatures. One of the important factors involved in the protective action of carotenoids is their influence on the molecular dynamics of membranes. To obtain complex information about interactions between carotenoids and lipids in a membrane, different techniques were used. In this review, the data resulting from EPR–spin label spectrometry, ³¹P- and ¹³C-NMR, differential scanning calorimetry, and computer simulation of the membrane molecular dynamics are presented. The effects of selected, structurally different carotenoid species on various physical parameters of model and natural membranes are described and their relevance to protection against some environmental stresses are discussed.     

Guard cell zeaxanthin tracks photosynthetically active radiation and stomatal apertures in Vicia faba leaves*

Zeaxanthin, antheraxanthin and violaxanthin concentrations in guard cells from sonicated abaxial epidermal peels of Vicia faba were measured from dawn to dusk, and compared with concentrations in mesophyll tissue of the same leaves. Measured changes in guard cell zeaxanthin and violaxanthin concentrations indicate that guard cells operate the xanthophyll cycle throughout the day. Mesophyll tissue had no detectable zeaxanthin at dawn, whereas guard cells had 30–50 mmol mol^?1 chlorophyll a+b. On a chlorophyll basis, maximal zeaxanthin levels were 3–4 fold higher in guard cells than in mesophyll cells. Zeaxanthin concentrations tracked levels of photosynthetically active radiation (PAR) in both mesophyll and guard cells. In the mesophyll, most of the zeaxanthin changes occurred in mid-morning and mid-afternoon. In guard cells, zeaxanthin concentrations changed nearly linearly with PAR in the early morning and late afternoon, and closely tracked PAR levels throughout the day. Guard cell zeaxanthin concentrations were also closely correlated with stomatal apertures. The close relationship between zeaxanthin concentrations and PAR levels in guard cells indicates that zeaxanthin is well suited to function as a molecular photosensor in stomatal movements.     

Coupled response of stomatal and mesophyll conductance to light enhances photosynthesis of shade leaves under sunflecks

Light gradients within tree canopies play a major role in the distribution of plant resources that define the photosynthetic capacity of sun and shade leaves. However, the biochemical and diffusional constraints on gas exchange in sun and shade leaves in response to light remain poorly quantified, but critical for predicting canopy carbon and water exchange. To investigate the CO₂ diffusion pathway of sun and shade leaves, leaf gas exchange was coupled with concurrent measurements of carbon isotope discrimination to measure net leaf photosynthesis (A_n), stomatal conductance (g_s) and mesophyll conductance (g_m) in Eucalyptus tereticornis trees grown in climate controlled whole‐tree chambers. Compared to sun leaves, shade leaves had lower A_n, g_m, leaf nitrogen and photosynthetic capacity (A_max) but g_s was similar. When light intensity was temporarily increased for shade leaves to match that of sun leaves, both g_s and g_m increased, and A_n increased to values greater than sun leaves. We show that dynamic physiological responses of shade leaves to altered light environments have implications for up‐scaling leaf level measurements and predicting whole canopy carbon gain. Despite exhibiting reduced photosynthetic capacity, the rapid up‐regulation of g_m with increased light enables shade leaves to respond quickly to sunflecks.     

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.