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, β-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.     

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.     

The photosynthetic pathway of the roots of twelve epiphytic orchids with CAM leaves

The photosynthetic pathway of the roots (both the white velamentous main portions and the green, nonvelamentous tips) was investigated in twelve taxa (natural species and intergeneric hybrid cultivars) of epiphytic orchids having CAM leaves. All organs contained chlorophyll, and the a/b ratios indicate that the organs, especially the roots, are likely shade-adapted. Stable carbon isotope ratios of the tissues were near −15‰ for all organs, a value typical of obligate (constitutive) CAM plants. Values for root tissues were slightly lower (more negative) than those of the leaves. The presence of CAM in the leaves of these orchids did not ensure that their roots performed CAM photosynthesis. Further work is needed to address the questions raised in this study and to determine if the photosynthetic roots of these taxa are capable of assimilating atmospheric CO₂.     

Twig and foliar nutrient concentrations in relation to nitrogen and phosphorus supply in a eucalypt (Eucalyptus diversicolor F. Muell.) and an understorey legume (Bossiaea laidlawiana Tovey and Morris)

The sensitivity of tissue nutrient concentrations to changes in plant age and the supply of P and N was compared between leaves and associated twigs in two forest species. In a young regrowth stand, tissues were sampled on three occasions from the mid-crown position of karri (Eucalyptus diversicolor F. Muell.) and Bossiaea laidlawiana Tovey and Morris, a major understorey legume. Leaves and twigs were also sampled from young plants of B. laidlawiana growing in a mature eucalypt stand to which P treatments had been applied.Nitrogen application increased N concentrations in twigs of karri and B. laidlawiana, but not in leaves. Phosphorus application increased P concentrations in both leaves and twigs of karri but the average increases were proportionally greater in twigs (65%) than in leaves (36%). Over the sampling period, P concentrations in leaves declined, while those in twigs were relatively stable. In B. laidlawiana, P supply also had a larger effect on P concentrations in twigs than in leaves. Addition of 200 kg P ha^–1 increased average P concentrations in twigs by 109% in the regrowth stand and by 215% in the mature stand while the corresponding increases in leaves were only 11% and 27%. Concentrations of other nutrients in both species were also affected by N and P application, the most notable being a decline in the concentrations of the minor nutrients, Zn and Cu, with increased P supply.The increased N concentrations in twigs of karri, and the increased P concentrations in tissues of both species, were associated with responses of karri to added N and P, and of B. laidlawiana to added P. This indicates that tissue concentrations of N and P were generally below critical concentrations where N and P were not applied. The results show that for these species twigs may be a better tissue than leaves for diagnosing deficiencies or predicting N and P requirements. The ratio of P concentrations in twigs to P concentrations in leaves also increased with added P. It is suggested that this ratio may be a useful index if it reduces the variability caused by sampling position within the crown or genetic differences between plants.     

CONTRASTING PHOTOSYNTHETIC BEHAVIOR IN LEAVES AND TWIGS OF HYMENOCLEA SALSOLA,A GREEN-TWIGGED WARM DESERT SHRUB

The photosynthetic behavior of leaves and twigs was compared in Hymenoclea salsola T. and G., a subshrub of the Mohave and Sonoran deserts, in which both leaves and green twigs make substantial contributions to whole-plant carbon gain. Light saturated photosynthesis in twigs was 0.62 times that of leaves (36.9 μmol m^-2 s^-1) when plants were well watered. Similar ratios were consistently observed in contrasting the photosynthetic responses of the two organ types to light, temperature, and intercellular CO₂, regardless of whether rates were compared under saturating or highly limiting conditions of light or intercellular CO₂. These scalar differences in photosynthetic rate between leaves and green twigs under a wide range of conditions were correlated with contrasting anatomical features such as chlorenchyma volume per projected area. Under normal ambient CO₂ concentrations (350 μl 1^-1), twigs on well watered plants operated at lower intercellular CO₂ concentrations than the leaves. Possible causes of this difference are discussed with respect to performance under well-watered conditions, organ lifespans, and contrasting anatomical constraints. Twigs require larger investments than do leaves of both carbon and nitrogen per projected area of the respective organs, yet they realize lower photosynthetic rates per intercepted light. Twigs, however, fulfill additional roles besides photosynthesis such as structural support and vascular transport which does not allow them to be as anatomically specialized as leaves for photosynthesis. Twigs also have a longer expected lifespan than leaves with a larger fraction of them surviving the summer drought period. This was correlated with a greater tolerance of twig than leaf photosynthesis to low plant water potentials.     

On the role of orientation in reducing photoinhibitory damage in photosynthetic-twig desert shrubs

Quantum yields for O₂ evolution were measured in photosynthetic twigs and leaves of Hymenoclea salsola and Senecio douglasii, two common shrubs of the deserts of western North America. When exposed to long-term drought, quantum yields of leaves and twigs remained constant up to the point of leaf abscision in both species. As water stress developed further, quantum yields began to decline in twigs; the extent of this quantum yield reduction was dependent on incident photon flux density. The reduction in quantum yield in twigs, which have a near-vertical orientation, was greatly accelerated when twigs were reoriented to the near-horizontal inclination typical of leaves. The reductions in quantum yield were not rapidly reversible and are interpreted as indicating photoinhibitory damage. The results are discussed in terms of the role that a near-vertical orientation might serve in maintaining photosynthetic structures through a drought period.     

Leaf carbon isotope and mineral composition in subtropical plants along an irradiance cline

Summary Leaf carbon isotope ratios and leaf mineral composition (Ca, K, Mg, Mn, N, and P) were measured on the dominant species along an irradiance cline in a subtropical monsoon forest of southern China. This irradiance cline resulted from disturbance caused by fuel-harvesting. Leaf carbon isotope ratios increased from undisturbed to disturbed sites for all species, indicating that leaf intercellular CO₂ concentrations decreased and leaf water use efficiencies increased along this cline. Nitrogen and magnesium levels were lower in leaves of species on the disturbed sites, but there were no clear patterns for calcium, potassium, phosphorus or manganese.C.I.W.D.P.B. Publication no 932     

Photosynthetic pigment contents in twigs of 24 woody species assessed by in vivo reflectance spectroscopy indicate low chlorophyll levels but high carotenoid/chlorophyll ratios

We have examined whether spectral reflectance indices used to non-destructively assess photosynthetic pigment levels and their ratios in leaves, could also be used for the same purpose in peridermal twigs. Regression lines of selected indices versus actual pigment levels, obtained from leaves and twigs of five species, suggested that semi-quantitative assessments are safe, provided that twig periderms could be easily removed. Given that, we proceeded to our next objective of screening a large number of species (24), in order to characterize their photosynthetic pigment profiles. Index comparisons between twigs and corresponding leaves indicated that twigs are characterized by lower levels of total chlorophyll and, unexpectedly, higher carotenoid/chlorophyll ratios. Moreover, the exposed and shaded sides of twigs displayed similar values for both indices in 80% of the species, suggesting that shade may not be the only factor shaping pigment levels and ratios. We discuss our results arguing that the distinct microenvironment within a twig may pose additional needs to the photosynthetic machinery, necessitating elevated carotenoid/chlorophyll ratios.     

Gender differences in Salix myrsinifolia at the pre‐reproductive stage are little affected by simulated climatic change

Females of dioecious species are known often to prioritize defense, while males grow faster. As climatic change is known to influence both growth and defense in plants, it would be important to know whether it affects the sexes of dioecious species differently. This could have impacts on future sex ratios in nature. We grew four clones of each sex of Salix myrsinifolia in greenhouse chambers under ambient conditions, enhanced temperature, enhanced CO₂ or enhanced temperature + enhanced CO₂. The females had the greatest growth and also the highest levels of phenolic compounds in twigs, while in leaves some compounds were higher in males, some in females. Enhanced CO₂ increased growth equally in both sexes, while growth was not affected by elevated temperature. Phenolic compounds in twigs were, however, lowered under elevated temperature. The gender differences were not strongly affected by the simulated climatic changes, but the effects seen on some highly concentrated compounds may be important. We interpret the intensive growth at pre‐reproductive phase as a strategy in females to get an initial advantage before later periods with fewer resources available for growth.     

Does elevated atmospheric CO2 concentrations affect wood decomposition?

This study was conducted to test the hypothesis that wood tissues generated under elevated atmospheric [CO₂] have lower quality and subsequent reduced decomposition rates. Chemical composition and subsequent field decomposition rates were studied for beech (Fagus sylvatica L.) twigs grown under ambient and elevated [CO₂] in open top chambers. Elevated [CO₂] significantly affected the chemical composition of beech twigs, which had 38% lower N and 12% lower lignin concentrations than twigs grown under ambient [CO₂]. The strong decrease in N concentration resulted in a significant increase in the C/N and lignin/N ratios of the beech wood grown at elevated [CO₂]. However, the elevated [CO₂] treatment did not reduce the decomposition rates of twigs, neither were the dynamics of N and lignin in the decomposing beech wood affected by the [CO₂] treatment, despite initial changes in N and lignin concentrations between the ambient and elevated [CO₂] beech wood. This revised version was published online in June 2006 with corrections to the Cover Date.     

Phytoextraction capacity of trees growing on a metal contaminated soil

Phytoremediation is an innovative biological technique to reclaim land contaminated by heavy metals or organic pollutants. In the present work, we studied the ability of five woody species to extract heavy metal (copper, zinc or cadmium) from a polluted soil to their above-ground tissues. Metal content in leaves and twigs was determined. Salix and Betula transferred zinc and cadmium to leaves and twigs, but Alnus, Fraxinus and Sorbus excluded them from their above-ground tissues. None of the species considered transferred copper to the shoots.     

Colonization of Japanese beech leaves by phyllosphere fungi

To examine the colonization of living leaves from buds and twigs by phyllosphere fungi of Japanese beech, the mycobiota were investigated on buds and twigs and on leaves covered with well-ventilated bags before budbreak. Ten phyllosphere fungi were isolated from rolled-up leaves within buds, bud scales, and twigs. However, frequencies of phyllosphere fungi on expanded leaves were reduced markedly when the buds were covered with well-ventilated bags before budbreak compared to the leaves that were not covered. This observation suggests that invasion of the fungi to the leaves from buds and twigs may be possible but is not the main route. Horizontal transmission may be common in endophytes and epiphytes of beech leaves. Phyllosphere mycobiota were then compared between sun and shade leaves. Of 13 species recorded as phyllosphere fungi, the frequencies of 2 species were lower and those of 3 species were higher in sun leaves than in shade leaves. Frequencies of the other 8 phyllosphere species were not different between sun and shade leaves. This result indicates that the colonization of leaves by some phyllosphere fungi was affected by the microenvironmental conditions on leaf surfaces.     

Short-term variation in the isotopic composition of organic matter allocated from the leaves to the stem of Pinus sylvestris: effects of photosynthetic and postphotosynthetic carbon isotope fractionation

We aimed to quantify the separate effects of photosynthetic and postphotosynthetic carbon isotope discrimination on δ¹³C of the fast‐turn‐over carbon pool (water soluble organic carbon and CO₂ emitted from heterotrophic tissues), including their diel variation, along the pathway of carbon transport from the foliage to the base of the stem. For that purpose, we determined δ¹³C in total and water‐soluble organic matter of the foliage plus δ¹³C and δ¹⁸O in phloem organic matter of twigs and at three heights along the stem of Pinus sylvestris over a nine‐day period, including four measurements per day. These data were related to meteorological and photosynthesis parameters and to the δ¹³C of stem‐emitted CO₂. In the canopy (foliage and twigs), the δ¹³C of soluble organic matter varied diurnally with amplitudes of up to 1.9‰. The greatest ¹³C enrichment was recorded during the night/early morning, indicating a strong influence of starch storage and remobilization on the carbon isotope signatures of sugars exported from the leaves. ¹³C enrichment of soluble organic matter from the leaves to the twig phloem and further on to the phloem of the stem was supposed to be a result of carbon isotope fractionation associated with metabolic processes in the source and sink tissues. CO₂ emitted from the stem was enriched by 2.3–5.2‰ compared with phloem organic matter. When day‐to‐day variation was addressed, water‐soluble leaf δ¹³C and twig phloem δ¹⁸O were strongly influenced by c_i/c_a and stomatal conductance (G_s), respectively. These results show that both photosynthetic and postphotosynthetic carbon isotope fractionation influence δ¹³C of organic matter over time, and over the length of the basipetal transport pathway. Clearly, these influences on the δ¹³C of respired CO₂ must be considered when using the latter for partitioning of ecosystem CO₂ fluxes or when the assessment of δ¹³C in organic matter is applied to estimate environmental effects in c_i/c_a.     

Age-dependent bark photosynthesis of aspen twigs

The photosynthetic performance of trembling aspen (Populus tremula L.) twigs and leaves was studied in relation to selected structural features of aspen bark. PFD transmittance of intact periderm was reduced by about 90% in current-year twigs through peridermal thickening. However, because of drastic changes within the bark microstructure, PFD transmittance increased in 1-year-old twig segments up to 26% of the incident PFD. On a unit surface area basis, the chlorophyll content of young twigs (425 mg Chl m^-2) almost reached that of leaves (460 mg Chl m^-2). The chlorophyll content of aspen bark chlorenchyma was clearly age-dependent, even increasing in current-year twigs with advancing internodal age. The low bark chlorophyll a/b ratios (about 2.6 compared with 3.9 in leaves) indicate that bark chloroplasts are shade-adapted. Positive net photosynthesis was not found in aspen twigs, but apparent respiration was distinctly reduced in the light due to light-driven carbon refixation (bark photosynthesis) within the chlorenchymal tissues. Under constant microclimatic conditions, dark respiration rates were strongly correlated with stem-internal CO₂ refixation. In accordance with increasing dark respiration rates, the efficiency of this carbon recycling was generally greater in the metabolically more active, younger twig segments than in older segments; carbon refixation rates reached up to 80% of dark respiration values. At least in young twigs and branches and thus in the light-exposed outer parts of tree crowns, respiratory CO₂ losses by the tree skeleton could efficiently be reduced. Refixation of carbon dioxide may be of great importance for carbon budgets in the environmentally controlled or pathogen-induced leafless states of deciduous aspen trees.     

Carbon, hydrogen, and oxygen isotope ratios of cellulose from plants having intermediary photosynthetic modes

Carbon and hydrogen isotope ratios of cellulose nitrate and oxygen isotope ratios of cellulose from species of greenhouse plants having different photosynthetic modes were determined. When hydrogen isotope ratios are plotted against carbon isotope ratios, four clusters of points are discernible, each representing different photosynthetic modes: C₃ plants, C₄ plants, CAM plants, and C₃ plants that can shift to CAM or show the phenomenon referred to as CAM-cycling. The combination of oxygen and carbon isotope ratios does not distinguish among the different photosynthetic modes. Analysis of the carbon and hydrogen isotope ratios of cellulose nitrate should prove useful for screening different photosynthetic modes in field specimens that grew near one another. This method will be particularly useful for detection of plants which show CAM-cycling.     

The effects of tissue type and body size on δ13C and δ15N values in parrotfish (Labridae) from Zanzibar,Tanzania

Differences between the stable isotopic ratios (δ¹³C and δ¹⁵N) of two tissues (blood and muscle) from four species of East African coral reef parrotfishes (family: Labridae, tribe: Scarini) were analysed across a broad spectrum of body sizes. Comparison of isotopic ratios between the tissues allowed the assessment of using blood as an alternative tissue to muscle. In 2010–2011, constant differences between tissues (δ_blood minus δ_muscle) were found across a broad range of sampled fish lengths. Linear relationships between the tissues, specific for an isotope, indicate that constants could be generated for converting blood isotope into muscle isotope values. Only one species, Chlorurus sordidus, displayed an inconsistent difference between tissues in δ¹⁵N, indicating that this ratio was dependent on fish length. The δ¹³C of both tissues was positively related linearly to fish length for three species, while δ¹⁵N showed no relationship with body length. The results are interpreted as indicating dietary consistency over days to weeks, the time of tissue turnover for blood and muscle, respectively. Lastly, differences among the species, even closely related species, show that the generation of tissue conversion constants is species‐specific.     

Two distinct plant respiratory physiotypes might exist which correspond to fast-growing and slow-growing species

The origin of the carbon atoms in CO₂ respired by leaves in the dark of several plant species has been studied using ¹³C/¹²C stable isotopes. This study was conducted using an open gas exchange system for isotope labeling that was coupled to an elemental analyzer and further linked to an isotope ratio mass spectrometer (EA–IRMS) or coupled to a gas chromatography–combustion-isotope ratio mass spectrometer (GC–C-IRMS). We demonstrate here that the carbon, which is recently assimilated during photosynthesis, accounts for nearly ca. 50% of the carbon in the CO₂ lost through dark respiration (R_d) after illumination in fast-growing and cultivated plants and trees and, accounts for only ca. 10% in slow-growing plants. Moreover, our study shows that fast-growing plants, which had the largest percentages of newly fixed carbon of leaf-respired CO₂, were also those with the largest shoot/root ratios, whereas slow-growing plants showed the lowest shoot/root values.     

Activity ratios of ribulose-1,5-bisphosphate carboxylase accurately reflect carbamylation ratios

Activity ratios and carbamylation ratios of ribulose-1,5-bisphosphate carboxylase (RuBPCase) were determined for leaves of Phaseolus vulgaris and Spinacia oleracea exposed to a variety of partial pressures of CO₂ and O₂ and photon flux densities (PFD). It was found that activity ratios accurately predicted carbamylation ratios except in extracts from leaves held in low PFD. In particular, it was confirmed that the loss of RuBPCase activity in low partial pressure of O₂ and high PFD results from reduced carbamylation. Activity ratios of RuBPCase were lower than carbamylation ratios for Phaseolus leaves sampled in low PFD, presumably because of the presence of 2-carboxyarabinitol 1-phosphate. Spinacia leaves sampled in darkness also exhibited lower activity ratios than carbamylation ratios indicating that this species may also have an RuBPCase inhibitor even though carboxyarabinitol 1-phosphate has not been detected in this species in the past.     

Effect of stem length to stem slender ratio of current-year twigs on the leaf display efficiency in evergreen and deciduous broadleaved trees

Aims Branches and leaves are the two main structural units of tree crown composition. Among the adaptive strategies of plants, the functional traits of branches and the relationships between branch traits and leaf traits determine the capacity of trees to access light and space. In this study, our objective is to test the hypothesis that leaf display efficiency is affected by the stem length to stem slender ratio within current-year twigs.Methods The stem length to stem slender ratios of current-year twigs were used as the proxy of stem structure traits. Leaf area ratio (total leaf area per stem mass), leaf density (leaf number per stem length) and leaf/stem mass ratio (total leaf mass per stem mass) were used as the proxies of leaf display efficiency. The relationship between stem structure traits and leaf display efficiency within current-year twigs were studied for 25 evergreen and 60 deciduous broadleaved woody species in Qingliang Mountain, Zhejiang, China. The standardized major axis estimation method was used to examine the scaling relationship between stem structural traits and leaf display efficiency within current-year twigs.Important findings The proxies of leaf display efficiency, measured by leaf area ratio, leaf density or leaf/stem mass ratio, were all significantly and negative correlated with stem length to stem slender ratio within current-year twigs in both evergreen and deciduous broadleaved woody species. This suggested that leaf display efficiency decreased with stem length to stem slender ratios within current-year twigs, which may reflect the role of mechanical safety and light within twigs. The slope of the relationship between leaf display efficiency and stem long-dimension structure traits in evergreen species was not significantly different from the one in deciduous species. In contrast, the y-intercept of the relationship between leaf density and stem long-dimension structure traits was significantly larger in evergreen species than in deciduous species, i.e. the leafing intensity of evergreen species was higher than that of deciduous species. Individual leaf area and specific leaf area were smaller in evergreen species than in deciduous species, which resulted in deciduous species have a larger leaf area per stem mass and leaf mass per stem mass at a given stem length to stem slender ratio compared to evergreen species. It may reflect the conservative adaptive strategy of high consumption and slow benefit in evergreen species. Our results demonstrated that leaf display efficiency could be affected by stem length, and would change with leaf life-span (deciduous versus evergreen).     

Fungal Endophytic Communities in Healthy and Declining Quercus robur L. and Q. cerris L. Trees in Northern Italy

Healthy and declining English oak (Quercus robur) and Turkey oak (Q. cerris) in north‐western Italy, in a plain oak forest showing decline for oak puzzle disease, were tested to assess possible variations in the composition of their fungal endophytic communities and their relation to the health status of trees. Samples collected in spring (buds) and in autumn (leaves, annual shoots and twigs) were surface‐sterilized, cut into fragments and placed on potato dextrose agar for a month; 26 fungal species were isolated, the most frequent being Tubakia dryina, Dendrodochium sp., Eutypella sp. and a sterile mycelium. Correspondence analysis showed significant qualitative differences between assemblages inhabiting twigs and herbaceous tissues that were due to the low frequency of Tubakia dryina in twigs and its higher frequency in buds, leaves and shoots. Tubakia dryina was isolated more frequently from leaves of declining oaks and from buds of healthy oaks; Monochaetia monochaeta showed a preference for healthy trees, especially leaves and buds. According to the Shannon–Wiener index, endophytic fungal communities of leaves, twigs and buds of declining English oak were poorer than those of declining Turkey oak, but there were no significant differences between healthy hosts.