Mineral cost of carnivory in aquatic carnivorous plants

Tissue N, P, K, Ca, and Mg content was estimated in traps and photosynthetic and carnivorous shoots in five aquatic carnivorous plant species from an outdoor culture: Aldrovanda vesiculosa, Utricularia vulgaris, U. reflexa, U. intermedia, and U. stygia, for the determination of the mineral cost of carnivory. In three species with monomorphic shoots (A. vesiculosa, U. vulgaris, U. reflexa), tissue P and K content in traps was significantly higher than that in their photosynthetic shoots, whereas N content was about the same. In U. stygia and U. intermedia with dimorphic shoots, tissue N and P content was markedly the highest in photosynthetic shoots followed by traps, while it was lowest in carnivorous shoots. In all five species, trap K content was significantly (2–4 times) higher than that in photosynthetic and carnivorous shoots. In all species, the values of the mineral cost of carnivory – the proportion of mineral nutrient amount contained in traps or carnivorous shoots to that in the total plant biomass – were within 19–61% for N, 33–76% P, 51–78% K, 26–70% Ca, and 34% for Mg. A new concept of the ecological cost-benefit relationships of plant carnivory, based on the mineral benefit of prey capture and mineral costs associated with trap production, is introduced for aquatic carnivorous plants. The evolution of this plant group is considered to show the optimization of these mineral cost-benefit relationships.     

Variation of sterility and fertility in Utricularia australis f. australis in Hokkaido, northern Japan

Seed-production ability was studied in Utricularia australis R. Br. f. australis Komiya and Shibata through field observation, pollination experiment, pollen culture, and isozyme analysis for populations in various regions of Hokkaido, northern Japan. Utricularia australis f. australis has previously been postulated to be sterile in Japan, however, in the present study, fertile populations were found in eastern Hokkaido. Bisexual sterility and male sterility were suspected to be present in populations in the western part of Hokkaido. There was also a strain that seemed to produce no seed because of intra-clonal inbreeding depression or self-incompatibility. Isozyme analysis of three enzyme systems indicated a uniformity of multienzymatic phenotype (MEP) in each population or region. The MEP grouping of populations corresponded to the type of sterility and fertility revealed by pollination and pollen culture experiments. These experiments suggested that each regional strain had its own sterility or fertility type.     

The influence of prey capture on photosynthetic rate in two aquatic carnivorous plant species

Photosynthetic (P_N) and dark respiration rate (RD) were measured in two species of aquatic carnivorous plants, Aldrovanda vesiculosa and Utricularia australis, growing with or without prey in an outdoor growth experiment. After 7–14 days, the positive growth effect of feeding on prey (apical growth rate, plant size, branching) was evident in both species. Tissue N content in young leaf whorls of both species fed on prey was significantly lower (by 54% in Aldrovanda, by 86% in Utricularia) than that in unfed variants, while tissue P content was about the same. In both species, chlorophyll a content was consistent with tissue N content and RD in fed and unfed variants was very similar. However, P_N was significantly greater in fed plants of Aldrovanda, while lower in Utricularia. It is possible to conclude that feeding on prey in aquatic carnivorous plants leads to neither an increase of shoot tissue N content nor P_N per unit biomass. Thus, the main physiological effect of catching prey is not enhancement of P_N, but to provide N and P for essential growth processes in juvenile tissues.     

Oscillations of the internal CO(2) concentration in tobacco leaves transferred to low CO(2)

Measurement of the internal CO(2) concentration (Ci) in tobacco leaves using a fast-response CO(2) exchange system showed that in the light, switching from 350 microLL(-1) to a low CO(2) concentration of 36.5 microLL(-1) (promoting high photorespiration) resulted in the Ci oscillating near the value of CO(2) compensation point (Gamma*). The oscillations are highly irregular, the range of Ci varying by 2-4 microLL(-1) in substomatal cavities with a period of a few seconds. The statistical properties of the time series became stationary after a transient of approximately 100s following transfer to low CO(2). Attractor reconstruction shows that the observed oscillations are not chaotic but exhibit stochastic behavior. The period of oscillations is consistent with the duration of photorespiratory post-illumination burst (PIB). We suggest that the observed oscillations may be due to a similar mechanism to that which leads to PIB, and may play a role in switching mitochondrial operation between oxidation of the photorespiratory glycine and of the tricarboxylic acid cycle substrates.     

Restricted seed contribution and clonal dominance in a free-floating aquatic plant <Emphasis Type="Italic">Utricularia australis</Emphasis> R. Br. in southwestern Japan

Utricularia australis R. Br. is an aquatic angiosperm species common in natural and irrigation ponds in temperate regions. This species reproduces both sexually and vegetatively, but in southwestern Japan the occurrence of male-sterile populations, in which plants produce no pollen and propagate only vegetatively, has been recorded. We studied the reproductive contribution of seeds in normal pollen-producing populations using isozyme analyses, a pollination experiment under culture and field observations. Although seedlings obtained from controlled mating indicated segregation of isozyme, polymorphism of the isozyme genotype was detected mainly among populations, but rarely within each pond population. This suggested clonal dominance and rarity of seed or seedling survival in natural populations. In the pollination experiment, the mean seed set ratio in cross-pollination between plants of the same isozyme genotype (7.6%) did not differ significantly from self-pollination (7.6%), but was lower than cross-pollination between plants of different genotypes (45.7%). The low ratio in crossing between the same genotype plants was ascribed to the clonality of the parents. In general, these results corresponded with the low ratios in seed setting observed in natural populations (7.9–13.7%). All the male-sterile populations we surveyed showed the same genotype, thus male sterility in the study area was considered to have the same origin.     

Photosynthetic characteristics of coffee (Coffea arabusta) plantlets in vitro in response to different CO2 concentrations and light intensities

The photosynthetic characteristics of coffee ( Coffea arabusta) plantlets cultured in vitro in response to different CO₂ concentrations inside the culture vessel and photosynthetic photon flux (PPF) were investigated preliminarily. The estimation of net photosynthetic rate (P_n) of coffee plantlets involved three methods: (1) estimating time courses of actual P_n in situ based on measuring CO₂ concentrations inside and outside the vessel during a 45-day period, (2) estimating P_n in situ at different CO₂ concentrations and PPFs using the above measuring approach for 10-day and 30-day old in vitro plantlets, and (3) estimating P_n of a single leaf at different CO₂ concentrations and PPFs by using a portable photosynthesis measurement system for 45-day old in vitro coffee plantlets. The results showed that coffee plantlets in vitro had relatively high photosynthetic ability and that the P_n increased with the increase in CO₂ concentration inside the vessel. The CO₂ saturation point of in vitro coffee plantlets was high (4500–5000 μmol mol^-1); on the other hand, the PPF saturation point was not so high as compared to some other species, though it increased with increasing CO₂ concentration inside the vessel. This revised version was published online in June 2006 with corrections to the Cover Date.     

Land plants equilibrate O2 and CO2 concentrations in the atmosphere

The role of land plants in establishing our present day atmosphere is analysed. Before the evolution of land plants, photosynthesis by marine and fresh water organisms was not intensive enough to deplete CO₂ from the atmosphere, the concentration of which was more than the order of magnitude higher than present. With the appearance of land plants, the exudation of organic acids by roots, following respiratory and photorespiratory metabolism, led to phosphate weathering from rocks thus increasing aquatic productivity. Weathering also replaced silicates by carbonates, thus decreasing the atmospheric CO₂ concentration. As a result of both intensive photosynthesis and weathering, CO₂ was depleted from the atmosphere down to low values approaching the compensation point of land plants. During the same time period, the atmospheric O₂ concentration increased to maximum levels about 300 million years ago (Permo-Carboniferous boundary), establishing an O₂/CO₂ ratio above 1000. At this point, land plant productivity and weathering strongly decreased, exerting negative feedback on aquatic productivity. Increased CO₂ concentrations were triggered by asteroid impacts and volcanic activity and in the Mesozoic era could be related to the gymnosperm flora with lower metabolic and weathering rates. A high O₂/CO₂ ratio is metabolically linked to the formation of citrate and oxalate, the main factors causing weathering, and to the production of reactive oxygen species, which triggered mutations and stimulated the evolution of land plants. The development of angiosperms resulted in a decrease in CO₂ concentration during the Cenozoic era, which finally led to the glacial-interglacial oscillations in the Pleistocene epoch. Photorespiration, the rate of which is directly related to the O₂/CO₂ ratio, due to the dual function of Rubisco, may be an important mechanism in maintaining the limits of O₂ and CO₂ concentrations by restricting land plant productivity and weathering.     

The Photosynthetic Response to Elevated CO2 in High Altitude Potato Species (Solanum curtilobum)

Plants of Solanum curtilobum (from high altitude) and Solanum tuberosum (from low altitude) were grown in open-top chambers in a greenhouse at either ambient (AC, 360 µmol mol^–1) or ca. twice ambient (EC, 720 µmol mol^–1) CO₂ concentrations for 30 d. CO₂ treatments started at the reproductive stage of the plants. There were similar patterns in the physiological response to CO₂ enrichment in the two species. Stomatal conductance was reduced by 59 % in S. tuberosum and by 55 % in S. curtilobum, but such a reduction did not limit the net photosynthetic rate (P _N), which was increased by approximately 56 % in S. curtilobum and 53 % in S. tuberosum. The transpiration rate was reduced by 16 % in both potato species while instantaneous transpiration efficiency increased by 80 % in S. tuberosum and 90 % in S. curtilobum. Plants grown under EC showed 36 and 66 % increment in total dry biomass, whereas yields (dry mass of tubers) were increased by 40 and 85 % in S. tuberosum and S. curtilobum, respectively. EC promoted productivity by increasing P _N. Thus S. tuberosum, cultivated around the world at low altitudes, and S. curtilobum, endemic of the highland Andes, respond positively to EC during the tuberisation stage.     

Capture of algae promotes growth and propagation in aquatic Utricularia

Background and Aims Some carnivorous plants trap not only small animals but also algae and pollen grains. However, it remains unclear if these trapped particles are useless bycatch or whether they provide nutrients for the plant. The present study examines this question in Utricularia, which forms the largest and most widely spread genus of carnivorous plants, and which captures prey by means of sophisticated suction traps.Methods Utricularia plants of three different species (U. australis, U. vulgaris and U. minor) were collected in eight different water bodies including peat bogs, lakes and artificial ponds in three regions of Austria. The prey spectrum of each population was analysed qualitatively and quantitatively, and correlated with data on growth and propagation, C/N ratio and δ¹⁵N.Key Results More than 50 % of the prey of the Utricularia populations investigated consisted of algae and pollen, and U. vulgaris in particular was found to capture large amounts of gymnosperm pollen. The capture of algae and pollen grains was strongly correlated with most growth parameters, including weight, length, budding and elongation of internodes. The C/N ratio, however, was less well correlated. Other prey, such as moss leaflets, fungal hyphae and mineral particles, were negatively correlated with most growth parameters. δ¹⁵N was positively correlated with prey capture, but in situations where algae were the main prey objects it was found that the standard formula for calculation of prey-derived N was no longer applicable.Conclusions The mass capture of immotile particles confirms the ecological importance of autonomous firing of the traps. Although the C/N ratio was little influenced by algae, they clearly provide other nutrients, possibly including phosphorus and trace elements. By contrast, mosses, fungi and mineral particles appear to be useless bycatch. Correlations with chemical parameters indicate that Utricularia benefits from nutrient-rich waters by uptake of inorganic nutrients from the water, by the production of more traps per unit of shoot length, and by the capture of more prey particles per trap, as nutrient-rich waters harbour more prey organisms.     

Decline in gypsy moth (Lymantria dispar) performance in an elevated CO2 atmosphere depends upon host plant species

Plant species differ broadly in their responses to an elevated CO₂ atmosphere, particularly in the extent of nitrogen dilution of leaf tissue. Insect herbivores are often limited by the availability of nutrients, such as nitrogen, in their host plant tissue and may therefore respond differentially on different plant species grown in CO₂-enriched environments. We reared gyspy moth larvae (Lymantria dispar) in situ on seedlings of yellow birch (Betula allegheniensis) and gray birch (B. populifolia) grown in an ambient (350 ppm) or elevated (700 ppm) CO₂ atmosphere to test whether larval responses in the elevated CO₂ atmosphere were species-dependent. We report that female gypsy moths (Lymantria dispar) reared on gray birch (Betula populifolia) achieved similar pupal masses on plants grown at an ambient or an elevated CO₂ concentration. However, on yellow birch (B. allegheniensis), female pupal mass was 38% smaller on plants in the elevated-CO₂ atmosphere. Larval mortality was significantly higher on yellow birch than gray birch, but did not differ between the CO₂ treatments. Relative growth rate declined more in the elevated CO₂ atmosphere for larvae on yellow birch than for those on gray birch. In preference tests, larvae preferred ambient over elevated CO₂-grown leaves of yellow birch, but showed no preference between gray birch leaves from the two CO₂ atmospheres. This differential response of gypsy moths to their host species corresponded to a greater decline in leaf nutritional quality in the elevated CO₂ atmosphere in yellow birch than in gray birch. Leaf nitrogen content of yellow birch dropped from 2.68% to 1.99% while that of gray birch leaves only declined from 3.23% to 2.63%. Meanwhile, leaf condensed tannin concentration increased from 8.92% to 11.45% in yellow birch leaves while gray birch leaves only increased from 10.72% to 12.34%. Thus the declines in larval performance in a future atmosphere may be substantial and host-species-specific.     

Effect at the ecosystem level of elevated atmospheric CO2in an aquatic microcosm

We studied the responses of an aquatic microcosm in two different eutrophic conditions to elevated atmospheric CO₂concentration. We used microcosms, consisting of Escherichia coli(bacteria), Tetrahymena thermophila(protozoa) and Euglena gracilis(algae), in salt solution with 50 and 500 mg l^–1of proteose peptone (eutrophic and hypereutrophic conditions, respectively) under ambient and elevated CO₂(1550±100 l l^–1) conditions. The density of E. gracilisincreased significantly under elevated CO₂in both eutrophic and hypereutrophic microcosms. In the eutrophic microcosm, the other elements were not affected by elevated CO₂. In the hypereutrophic microcosm, however, the concentrations of ammonium and phosphate decreased significantly under elevated CO₂. Furthermore, the density of T. thermophilawas maintained in higher level than that in the microcosm with ambient CO₂and the density of E. coliwas decreased by CO₂enrichment. Calculating the carbon biomasses of T. thermophilaand E. colifrom their densities, the changes in their biomasses by CO₂enrichment were little as compared with large increase of E. graciliscarbon biomass converted from chlorophyll a. From the responses to elevated CO₂in the subsystems of the hypereutrophic microcosm consisting of either one or two species, the increase of E. graciliswas a direct effect of elevated CO₂, whereas the changes in the density of E. coliand T. thermophilaand the decreases in the concentration of ammonium and phosphate are considered to be indirect effects rather than direct effects of elevated CO₂. The indirect effects of elevated CO₂were prominent in the hypereutrophic microcosm.     

Two new species of the carnivorous genusPinguicula, (Lentibulariaceae) from Mediterranean habitats

Two species ofPinguicula (P. submediterranea andP. mundi) are newly described from the south and east of the Iberian Peninsula. We analysed morphometric, cytogenetic, RAPD, ecological, and breeding system data to support the distinctness of these taxa.Pinguicula submediterranea andP. mundi are both hexaploid (2n = 48), a chromosome number previously not reported for the genus. Biometric, cytogenetical and molecular results distinguishP. submediterranea andP. mundi from otherPinguicula species, and from each other.Pinguicula mundi is predominantly outbreeding, whereasP. submediterranea both inbreeds and outbreeds.Pinguicula submediterranea andP. mundi appear to be isolated reproductively, although gene flow between the allopatricP. submediterranea andP. mundi is unlikely. BothP. submediterranea andP. mundi are threatened with extinction because suitable habitats are diminishing in size, or even disappearing, due to current aridity in the region.     

The CO2 sense of the moth Cactoblastis cactorum and its probable role in the biological control of the CAM plant Opuntia stricta

The interaction between the moth, Cactoblastis cactorum, and the cactus, Opuntia stricta, is used as a model to examine the question of whether the CO₂ sense of a herbivorous insect can detect the CO₂ gradients associated with a plant's metabolic activity. Both the anatomical and the electrophysiological characteristics of CO₂-sensitive receptor neurons in C. cactorum indicate an adaptation to the detection of small fluctuations around the atmospheric background. Evidence is provided that further rises in background will impair the function of the sensory organ. In the habitat of the plant, during the diurnal window of the moth's activity, two types of CO₂ gradients occur that are detectable by the moth's sensors. The first gradient, associated with soil respiration, is vertical and extends from the soil surface to an altitude of approximately 1 m. Its magnitude is well above the detectability limit of the sensors. The notion that this gradient provides, to a flying insect, a cue for the maintenance of a flight altitude favourable for host detection is supported by field observations of behaviour. The second gradient, associated with CO₂ fixation by the plant, extends from the surfaces of photosynthetic organs (cladodes) over a boundary layer distance of approximately 5 mm. Again, its magnitude is well above the detectability limit. The notion that this gradient provides, to a walking insect, a cue to the physiological condition of the plant is supported by the observation that females of C. cactorum, prior to oviposition, actively probe the plant surface with their CO₂ sensors. In a simulation of probing, pronounced responses of the sensors to the CO₂-fixing capacity of O. stricta are observed. We propose that by probing the boundary layer, females of C. cactorum can detect the healthiest, most active O. stricta cladodes, accounting for earlier observations that the most vigorous plants attract the greatest density of egg sticks.     

Effects of temperature,oxygen concentration,leaf age and seasonal variations on the CO2 compensation point of Lolium perenne L.

Various factors affect the CO₂ compensation point of detached leaves of Lolium perenne L. These include oxygen concentration, temperature, leaf age, and season (spring and summer). Analysis of the results using the model of G.D. Farquhar, S. von Caemmerer and J.A. Berry (1980) Planta 149, 78–90, indicates that some of the CO₂ evolved by leaves in the light is derived from sources other than photorespiration. It is suggested that the operation of the tricarboxylic acid in the light can account for most of this CO₂.Azcón-Bieto—experimental work was done in Barcelona, the further analyses white at the A.N.U.     

Response of NAD(P)H dehydrogenase complex to the alteration of CO2 concentration in the cyanobacterium Synechocystis PCC6803

An NADPH-specific NDH-1 sub-complex was separated by native-polyacrylamide gel electrophoresis and detected by activity staining from the whole cell extracts of Synechocystis PCC6803. Low CO2 caused an increase in the activity of this sub-complex quickly, accompanied by an evident increase in the expression of NdhK and PSI-driven NADPH oxidation activity that can reflect the activity of NDH-1-mediated cyclic electron transport. During incubation with high CO2, the activities of NDH-1 sub-complex and PSI-driven NADPH oxidation as well as the protein level of NdhK slightly increased at the beginning, but decreased evidently in various degrees along with incubation time. These results suggest that CO2 concentration in vitro as a signal can control the activity of NDH-1 complex, and NDH-1 complex may in turn function in the regulation of CO2 uptake.     

Increased CO2 and nutrient status changes affect phytomass and the production of plant defensive secondary chemicals in Salix myrsinifolia (Salisb.)

The effect of CO₂ enrichment (700 and 1050 ppm) on phytomass, soluble sugars, leaf nitrogen and secondary chemicals of three Salix myrsinifolia clones was studied in plants cultivated at very poor (sand seedlings) and moderate (peat seedlings) nutrient availability and under low illumination. The total shoot phytomass production of sand scedlings was less than 10% of that of the peat seedlings. Carbon dioxide increased the total shoot phytomass of peat seedlings. When the ambient carbon supply was doubled (to 700 ppm) the growth of sand seedlings was slightly enhanced but 1050 ppm CO₂ gave growth figures similar to those at the control CO₂ level. Leaf nitrogen content and total soluble sugar contents were significantly higher in peat seedlings than in sand seedlings. Leaf nitrogen showed a decreasing trend in relation to CO₂ increase. On the other hand, CO₂ did not have any clear-cut effect on total sugars. At the control CO₂ level the content of salicortin, which is a dynamic phenolic, was higher in the peat seedlings than in the sand seedlings, but salicin showed the opposite trend. CO₂ enrichment considerably decreased these phenolics in the peat seedlings. At the control CO₂ level, the content of more static phenolics, such as proanthocyanidins, was higher in sand seedlings. An increased carbon supply considerably increased static phenolics in the peat seedlings. Willow defence against generalist herbivores is moderately decreased by enhancement of atmospheric carbon dioxide.     

Effect of temperature on the CO2/O2 specificity of ribulose-1,5-bisphosphate carboxylase/oxygenase and the rate of respiration in the light

Responses of the rate of net CO₂ assimilation (A) to the intercellular partial pressure of CO₂ (p _i ) were measured on intact spinach (Spinacia oleracea L.) leaves at different irradiances. These responses were analysed to find the value of p _i at which the rate of photosynthetic CO₂ uptake equalled that of photorespiratory CO₂ evolution. At this CO₂ partial pressure (denoted ), net rate of CO₂ assimilation was negative, indicating that there was non-photorespiratory CO₂ evolution in the light. Hence was lower than the CO₂ compensation point, . Estimates of were obtained at leaf temperatures from 15 to 30°C, and the CO₂/O₂ specificity of ribulose 1,5-bisphosphate (RuBP) carboxylase/oxygenase (E.C. 4.1.1.39) was calculated from these data, taking into account changes in CO₂ and O₂ solubilities with temperature. The CO₂/O₂ specificity decreased with increasing temperature. Therefore we concluded that temperature effects on the ratio of photorespiration to photosynthesis were not solely the consequence of differential effects of temperature on the solubilities of CO₂ and O₂. Our estimates of the CO₂/O₂ specificity of RuBP carboxylase/oxygenase are compared with in-vitro measurements by other authors. The rate of nonphotorespiratory CO₂ evolution in the light (R _d ) was obtained from the value of A at . At this low CO₂ partial pressure, R _d was always less than the rate of CO₂ evolution in darkness and appeared to decrease with increasing irradiance. The decline was most marked up to about 100 mol quanta m^-2 s^-1 and less marked at higher irradiances. At one particular irradiance, however, R _d as a proportion of the rate of CO₂ evolution in darkness was similar in different leaves and this proportion was unaffected by leaf temperature or by [O₂] (ambient and greater). After conditions of high [CO₂] and high irradiance for several hours, the rate of CO₂ evolution in darkness increased and R _d also increased.Abbreviations and symbols A rate of net CO₂-assimilation - CO₂ compensation point - CO₂ compensation point in the absence of R _d - p _i intercellular partial pressure of CO₂ - R _d (day respiration) rate of non-photorespiratory CO₂ evolution in the light - R _n (night respiration) rate of CO₂ evolution in darkness - RuBP ribulose-1,5-bisphosphate - Rubisco RuBP carboxylase/oxygenase     

Involvement of Exported Photosynthetic Products in the CO2Exchange of the Skeletal Shoots of Pine

CO₂exchange in the leafy and skeletal parts of attached shoots of Pinus sylvestrisL. was measured with an infrared gas-analyzer in an open differential system during daylight hours. The ¹⁴CO₂assimilation rates in the leafy parts of shoots and ¹⁴CO₂evolution from current photosynthetic products in the lower skeletal part of shoots were measured in afternoons. Chlorophyll content was measured in the needles of the same shoot. The carbon of exported assimilates contributed only about 4% to CO₂exchange in the heterotrophic tree tissues. Only this component of CO₂evolution from the surface of the skeletal part of the tree was related to the losses of the net primary photosynthetic production (NPP) in the aboveground part of the pine stand during the current growth period.