ALBERT

Notes: The influence of vesicular–arbuscular mycorrhizal (M) colonization on biomass production and photosynthesis of Trifolium repens L. was investigated in two experiments in which the foliar nitrogen and phosphorus contents of non-mycorrhizal (NM) plants were manipulated to be no lower than that of M plants. Throughout both experiments there was a stimulation in the rate of CO2 assimilation of the youngest, fully expanded leaf of M compared with NM plants. In addition, M plants exhibited a higher specific leaf area compared with NM plants, a response that maximized the area available for CO2 assimilation per unit of carbon (C) invested. Despite the increased rate of photosynthesis in M plants there was no evidence that the additional C gained was converted to biomass production of M plants. It is suggested that this additional C gained by colonized plants was allocated to the mycorrhizal fungus and that it is the fungus, by acting as a sink for assimilates, that facilitated the stimulation in the rate of photosynthesis of the plant partner.

Notes: Dipterocarps dominate the canopy of lowland tropical rain forest in South-east Asia. Seedlings of these species form diverse assemblages on the forest floor where low irradiance severely limits their growth. Further growth depends largely upon the increased irradiance that can occur with the creation of canopy gaps. However, the response of dipterocarp seedlings to increased irradiance and their subsequent establishment in the canopy may be influenced by the availability of other resources, such as nutrient availability. We investigated the influence of nitrogen supply on aspects of the photosynthetic physiology and growth of seedlings of four dipterocarp species (Shorea leprosula, Shorea johorensis, Shorea oleosa and Dryobalanops lanceolata) growing under low irradiance, during transfer from low to high irradiance, and during subsequent growth at high irradiance. All four species increased growth and photosynthetic capacity in response to N-supply at high irradiances but not at low irradiance approximating that which can be expected to occur in the forest understorey. When seedlings grown at low irradiances and varying N-supply were exposed to a large increase in irradiance, all species showed some degree of initial photodamage (measured through chlorophyll fluorescence), the extent of which was similar between species but differed markedly depending on the pre-exposure growth irradiance and N-supply. Greater photodamage occurred in seedlings grown at lower compared with higher N-supply and irradiance. Despite these initial difference in the extent of this photodamage, all seedlings demonstrated a similar capacity to recover from damage. However, the alterations in the photosynthetic physiology of leaves during this recovery differed between species and depended on N-supply. Under high N-supply all species apart from S. oleosa increased photosynthetic capacity per unit chlorophyll following exposure to high irradiance by increasing photosynthetic capacity per unit leaf area while, under low N-supply, an increase in photosynthetic capacity per unit leaf only occurred in D. lanceolata. Our results suggest that variations in N-availability may have a much greater impact on the relative competitiveness of dipterocarp seedlings during the regenerative phase following canopy gap formation than physiological differences between seedlings. Our results demonstrate a potentially significant role for N-availability in the regeneration dynamics and distribution of canopy-dominating dipterocarp species.

Notes: A comparative analysis of daily carbon (C) budgets and aspects of the C physiology of clover (Trifolium repens L.) colonized by vesicular-arbuscular (VA) mycorrhizal fungi was carried out over a 70 d growth period under conditions designed to ensure that shoots of mycorrhizal (M) and non-mycorrhizal (NM) plants were of similar nutrient status. C budgets did not differ on day 24 but by day 42 M plants had a significantly higher rate of photosynthesis than their NM counterparts when expressed on a whole shoot basis or unit dry weight basis. As both sets of plants were of the same size it was concluded that this greater C gain was the result of increased sink strength provided by the mycorrhizal fungus. By day 53 M plants had become larger than their uncolonized counterparts and a sink-induced stimulation in the rate of photosynthesis was no longer apparent. M plants had higher root sucrose, glucose and fructose pools from day 24. Analyses suggested that these sugars were utilized for trehalose and lipid synthesis, for the production of the large extramatrical mycelium and for the support of the respiratory demands of the M root system. Increased C allocation to roots of M plants was associated with a stimulation of the activities of cell wall and cytoplasmic invertases and of sucrose synthase in roots colonized by VA fungi. Such increases in enzyme activity may provide the mechanism enabling increased partitioning of carbohydrate both to the M root system and the fungal symbiont.

Notes: Nitrogen assimilation was studied in the deciduous, perennial climber Clematis vitalba. When solely supplied with NO3– in a hydroponic system, growth and N-assimilation characteristics were similar to those reported for a range of other species. When solely supplied with NH4+, however, nitrate reductase (NR) activity dramatically increased in shoot tissue, and particularly leaf tissue, to up to three times the maximum level achieved in NO3– supplied plants. NO3– was not detected in plant material that had been solely supplied with NH4+, there was no NO3– contamination of the hydroponic system, and the NH4+-induced activity did not occur in tobacco or barley grown under similar conditions. Western Blot analysis revealed that the induction of NR activity, either by NO3– or NH4+, was matched by NR and nitrite reductase protein synthesis, but this was not the case for the ammonium assimilation enzyme glutamine synthetase. Exposure of leaf disks to N revealed that NO3– assimilation was induced in leaves directly by NO3– and NH4+ but not glutamine. Our results suggest that the NH4+-induced potential for NO3– assimilation occurs when externally sourced NH4+ is assimilated in the absence of any NO3– assimilation. These data show that the potential for nitrate assimilation in C. vitalba is induced by a nitrogenous compound in the absence of its substrate and suggest that NO3– assimilation in C. vitalba may have a significant role beyond the supply of reduced N for growth.

Notes: The reversible dissipation of excitation energy in higher plants is believed to protect against light-induced damage to the photosynthetic apparatus. This dissipation is measured as the non-photochemical quenching of chlorophyll fluorescence. A method is described whereby the saturated capacity for rapidly reversible non-photochemical quenching can be compared between plant species. This method was applied to 22 common British plant species whose habitat was quantified using an index that describes shade tolerance. An association was found between occurrence in open habitats and a high capacity for non-photochemical quenching. It was found that, whilst this capacity was species dependent, it did not depend upon the conditions under which the plant was grown. The possible role of zeaxanthin as a determinant of quenching capacity was examined by measuring the contents of xanthophyll cycle carotenoids for each species. Comparing species, no correlation was seen between the saturated level of non-photochemical quenching and zeaxanthin content expressed relative to either total carotenoid or to chlorophyll. When zeaxanthin was expressed relative to the amount of xanthophyll cycle intermediates (zeaxanthin, antheraxanthin and violaxanthin), a weak correlation was seen.

Notes: The survival of dipterocarp seedlings in the understorey of south-east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. Field measurements demonstrated that Shorea leprosula seedlings in a rain forest understorey received a high proportion of daily photon flux density at temperatures supra-optimal for photosynthesis (72% at ≥30 °C, 14% at ≥35 °C). To investigate the effect of high temperatures on photosynthesis during sunflecks, gas exchange and chlorophyll fluorescence measurements were made on seedlings grown in controlled environment conditions either, under uniform, saturating irradiance (approximately 539 µmol m−2 s−1) or, shade/fleck sequences (approximately 30 µmol m−2 s−1/approximately 525 µmol m−2 s−1) at two temperatures, 28 or 38 °C. The rate of light-saturated photosynthesis, under uniform irradiance, was inhibited by 40% at 38 °C compared with 28 °C. However, during the shade/fleck sequence, photosynthesis was inhibited by 59% at 38 °C compared with 28 °C. The greater inhibition of photosynthesis during the shade/fleck sequence, when compared with uniform irradiance, was driven by the lower efficiency of dynamic photosynthesis combined with lower steady-state rates of photosynthesis. These results suggest that, contrary to current dogma, sunfleck activity may not always result in significant carbon gain. This has important consequences for seedling regeneration processes in tropical forests as well as for leaves in other canopy positions where sunflecks make an important contribution to total photon flux density.

Notes: The survivorship of dipterocarp seedlings in the deeply shaded understorey of South-east Asian rain forests is limited by their ability to maintain a positive carbon balance. Photosynthesis during sunflecks is an important component of carbon gain. To investigate the effect of elevated CO2 upon photosynthesis and growth under sunflecks, seedlings of Shorealeprosula were grown in controlled environment conditions at ambient or elevated CO2. Equal total daily photon flux density (PFD) (∼7·7 mol m−2 d−1) was supplied as either uniform irradiance (∼170 µmol m−2 s−1) or shade/fleck sequences (∼30 µmol m−2 s−1/∼525 µmol m−2 s−1). Photosynthesis and growth were enhanced by elevated CO2 treatments but lower under flecked irradiance treatments. Acclimation of photosynthetic capacity occurred in response to elevated CO2 but not flecked irradiance. Importantly, the relative enhancement effects of elevated CO2 were greater under sunflecks (growth 60%, carbon gain 89%) compared with uniform irradiance (growth 25%, carbon gain 59%). This was driven by two factors: (1) greater efficiency of dynamic photosynthesis (photosynthetic induction gain and loss, post-irradiance gas exchange); and (2) photosynthetic enhancement being greatest at very low PFD. This allowed improved carbon gain during both clusters of lightflecks (73%) and intervening periods of deep shade (99%). The relatively greater enhancement of growth and photosynthesis at elevated CO2 under sunflecks has important potential consequences for seedling regeneration processes and hence forest structure and composition.

Notes: Orobanche species are holoparasites which are very efficient sinks for host-derived solutes. Here, we report the use of direct measurements of xylem sap solute concentrations and water fluxes, together with a modelling procedure to calculate element fluxes within an association between Orobanche cernua and its tobacco host. Infection of tobacco by the parasite markedly influenced carbon acquisition and partitioning; net fixation of carbon was 20% higher in infected tobacco compared with controls. Orobanche cernua caused a 84% increase in net carbon flux moving downward from the tobacco shoot and 73% of this carbon was intercepted by the parasite, almost entirely through the phloem (〉99%). Further, the parasite also exerted a large impact on the nitrogen relations of the plant, notably nitrate uptake was stimulated and the amino acid content of xylem sap was lower. The parasite also relied heavily on host phloem for the supply of other resources, with only 5 to 15% of N, and 16% of K, 23% of Na, 63% of Mg and 13% of S being derived from the xylem. Thus, we provide quantitative information on the phloem dependency of the parasite and show that host carbon and nitrogen metabolism is stimulated as a consequence of infection.

Notes: The achlorophyllous holoparasitic angiosperm Orobanche cernua reduced biomass accumulation of its tobacco host, such that 73-d-old plants achieved 29% of the biomass of control plants. The difference in biomass between infected and uninfected tobacco could be accounted for directly by diversion of dry matter to the parasite. Thus, the smaller infected plants were responsible for the production of as much dry matter in host and parasite as their larger uninfected counterparts. The productivity of the infected system was maintained by: (a) sustained production of leaf area (a greater leaf area ratio); (b) increased specific leaf area, and (c) delayed senescence. When tobacco was inoculated with different densities of the parasite, the amount of dry matter accumulated by the parasite was not changed, suggesting that a finite amount of resource was available to the parasite. The response of the host to infection can be explained by simple source–sink interactions and the data are discussed with respect to other parasitic angiosperm–host systems which show different types of responses.

Notes: 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.