Attraction of Culicoides brevitarsis Kieffer (Diptera: Ceratopogonidae) and Culex annulirostris Skuse (Diptera: Culicidae) to simulated visual and chemical stimuli from cattle

Abstract  Different host-seeking responses to cattle by Culicoides brevitarsis and Culex annulirostris were found using two-dimensional cattle-shapes and simulated stimuli. Culicoides brevitarsis was attracted to the respiratory chemicals CO₂ and CO₂ + octenol but not to octenol alone. It was attracted to cattle-shape when CO₂ + octenol was also present and when the shape was enhanced with visible and infrared light. Culex annulirostris was attracted to each chemical treatment but not to any visual stimuli. Orientation of the host, possibly in relation to wind direction, may also be important but requires further consideration. The detection of behavioural differences between the species was dependent on the positioning of attractants and traps in relation to the host image. Responses by C. brevitarsis to CO₂ + octenol and the basic shape were only recorded in sticky traps. Contact with the shape was recorded initially on the shape head but preference was shown later to be for the ridge-line of the back. The response by Cx. annulirostris to CO₂ + octenol was only recorded in light traps placed centrally and above the shape. It is proposed that the initial location of hosts by C. brevitarsis is by visual stimuli. Attraction to CO₂ + octenol is secondary and probably only occurs close to the host. The chemicals help the midge to recognise and come to the potential host where landing is induced visually near the interface between the backline of the host and the background. The primary response by Cx. annulirostris is to CO₂ and octenol although these did not appear to bring this insect on to the host. Covers could be used to protect cattle from C. brevitarsis (but not Cx. annulirostris ) by blocking the visual stimuli.     

Comparison of carbon dioxide, octenol and a host-odour as mosquito attractants in the Upper Rhine Valley, Germany

ield studies were conducted in the Upper Rhine Valley to determine the responses of mosquitoes to CDC traps baited with either CO₂, octenol, light or paired combinations of these. Among eight mosquito species caught, the attractant effect on trap catches was studied in the four most abundant: Aedes vexans, Ae.rossicus, Ae.cinereus and Culex pipiens .
Traps baited only with light or octenol caught few mosquitoes, whereas many were caught by traps baited with CO₂ alone or in combination with either of the other candidate attractants. CO₂ baited traps, with or without light, caught the most Aedes . The combination of CO₂ and octenol attracted most Cx pipiens , but this apparent synergy was not significant.
Using a caged hamster compared to CO₂ as bait in a CDC light-trap with only intermittent fan suction, the hamster attracted less mosquitoes than CO₂ emitted at a rate of 225 g/h on days 1 and 2, whereas on days 3 and 4 the smell from the hamster's cage became significantly more attractive than this rate of CO₂ for all species of mosquitoes.     

Differential responses of Aedes and Culex mosquitoes to octenol or light in combination with carbon dioxide in Queensland, Australia

Abstract. Field studies were conducted with EVS (encephalitis vector surveillance) traps in south-eastern Queensland, Australia, to determine the relative response rates of mosquitoes to three levels (0.1, 4.5 and 30mg/h) of 1-octen-3-ol (octenol) in combination with a standard bait of 2200 g carbon dioxide (CO₂), compared with CO₂ alone or CO₂ with light from a 6V incandescent bulb. Compared to CO₂ alone, Aedes vigilax collections increased significantly when CO₂ was supplemented by all three octenol emission levels, but not by the addition of light. Furthermore, the 4.5 and 30 mg/h release rate of octenol gave a significant increase in numbers of Ae.vigilax relative to that from CO₂+ light. In contrast, collections of Culex annulirostris and Culex sitiens were not enhanced significantly by either the addition of light or octenol at all three levels. Fewer Cx sitiens were collected with octenol released at 4.5 mg/h in comparison to CO₂ alone. These differential sampling rates should be taken into account when using EVS traps.     

Spatial feeding preferences of ornithophilic mosquitoes,blackflies and biting midges

The section of habitat used by particular bloodsucking insects when seeking bloodmeals may influence the spectrum of hosts to which they have access and consequently the diseases they transmit. The vertical distribution of ornithophilic bloodsucking Diptera (Culicidae, Simuliidae and Ceratopogonidae) was studied using bird‐baited traps set at both ground and tree canopy levels. In total, 1240 mosquito females of eight species, 1201 biting midge females of 11 species, and 218 blackfly females of two species were captured during 2003–2005. Culex pipiens (L.) (Diptera: Culicidae) was found to prefer ground‐level habitats, whereas Anopheles plumbeus (Stephens) (Diptera: Culicidae), biting midges [Culicoides spp. (Diptera: Ceratopogonidae)] and Eusimulium angustipes (Edwards) (Diptera: Simuliidae) preferred the canopy. The results of this study with regard to Cx. pipiens behaviour differ from those of most previous studies and may indicate different spatial feeding preferences in geographically separate populations. The occurrence of E. angustipes in the canopy is concordant with its role in the transmission of avian trypanosomes. These findings may be important for surveillance programmes focusing on ornithophilic Diptera which transmit various pathogenic agents.     

The role of Palmyra palm trees (Borassus flabellifer) and sand fly distribution in northeastern India

Visceral leishmaniasis (VL), known as Kala-azar in India, is a parasite transmitted by the bite of the sand fly vector Phlebotomus argentipes. Published information on the species indicates it is a poor flyer, mainly hopping and gliding. This study describes the vector as more arboreal than previously documented. Data collected indicate the ability of P. argentipes and Sergentomyia spp to attain vertical heights in Palmyra palm trees Borassus flabellifer up to 18.4 m above ground level. To determine if sand flies were either climbing the tree trunk to rest in the canopy or flying, sticky traps were set around the tree trunk and checked for captures overnight. CDC traps set in the palm tree canopy resulted in the capture of 5,067 sand flies, 3,990 of which were P. argentipes. Traps were set during daylight hours to determine if sand flies remained and rested in the canopy. A total of 128 sand flies were trapped over 29 trap days in the palm trees. With the CDC traps, 130 P. argentipes and no Sergentomyia spp were captured. The converse was true for the sticky traps set around tree trunks 3 m below the CDC traps. Of the 105 sand flies collected, only one was P. argentipes and 104 were Sergentomyia spp. As reported elsewhere, this indicates Sergentomyia spp tend to climb and hop, wheareas P. argentipes are capable of longer and more sustained flight. Data presented herein suggest that P. argentipes is more exophylic and exophagic than previously reported. These findings have implications for sand fly control.     

Elevated CO2 concentration, nitrogen use, and seed production in annual plants

Elevated atmospheric CO₂ concentration ([CO₂]) stimulates seed mass production in many species, but the extent of stimulation shows large variation among species. We examined (1) whether seed production is enhanced more in species with lower seed nitrogen concentrations, and (2) whether seed production is enhanced by elevated [CO₂] when the plant uses more N for seed production. We grew 11 annuals in open top chambers that have different [CO₂] conditions (ambient: 370 μmol mol⁻¹, elevated: 700 μmol mol⁻¹). Elevated [CO₂] significantly increased seed production in six out of 11 species with a large interspecific variation (0.84–2.12, elevated/ambient [CO₂]). Seed nitrogen concentration was not correlated with the enhancement of seed production by elevated [CO₂]. The enhancement of seed production was strongly correlated with the enhancement of seed nitrogen per plant caused by increased N acquisition during the reproductive period. In particular, legume species tended to acquire more N and produced more seeds at elevated [CO₂] than non-nitrogen fixing species. Elevated [CO₂] little affected seed [N] in all species. We conclude that seed production is limited primarily by nitrogen availability and will be enhanced by elevated [CO₂] only when the plant is able to increase nitrogen acquisition.     

Trapping oestrid parasites of reindeer: the response of Cephenemyia trompeand Hypoderma tarandi tobaited traps

Abstract. At 340–360 km North of the Arctic Circle in Norway, Hypoderma tarandi (L.) and Cephenemyia trompe (Modeer) females were caught in baited traps from 10 July to 21 August. During three summers, adverse climatic conditions inhibited flight activity of these oestrids on 56–68% of the days. Flies were not caught prior to or after these dates, nor at winds above 8 m/s, temperatures below 10C, light intensities below 20,000 lux, or during periods of rain or snow. C0₂-baited insect flight traps caught significantly more H. tarandi females than non-baited traps. However, neither a white reindeer hide or reindeer interdigital pheromone glands enhanced the attraction of C0₂to H. tarandi or C. trompe. Hypoderma tarandi females also were attracted to mobile people, but not to stationary individuals. There were no significant differences in the numbers of C.trompe or H.tarandi caught in C0₂-baited traps in a birch/willow woods, on the treeless vidda (=tundra-like biome), or at woodsrvidda ecotone sites. Flies were caught in traps on days when the nearest reindeer herds were 25–100 km away. Significantly more H.tarandi and C.trompe were caught from 09.30 to 14.30 hours than from 14.30 to 19.30 hours; no flies were caught from 20.00 to 07.00 hours (Norwegian Standard Time = NST). Because CO_z-baited traps caught from hundreds to thousands of mosquitoes, blackflies and Culicoides midges, when climatic conditions inhibited oestrid activity, reindeer aggregations and movements attributed to insect attacks during warm sunny days may be largely in response to attacks by H.tarandi and C.trompe.     

Stomatal responses to CO2 during a diel Crassulacean acid metabolism cycle in Kalanchoe daigremontiana and Kalanchoe pinnata

To investigate the diurnal variation of stomatal sensitivity to CO₂, stomatal response to a 30 min pulse of low CO₂ was measured four times during a 24 h time-course in two Crassulacean acid metabolism (CAM) species Kalanchoe daigremontiana and Kalanchoe pinnata , which vary in the degree of succulence, and hence, expression and commitment to CAM. In both species, stomata opened in response to a reduction in p CO₂ in the dark and in the latter half of the light period, and thus in CAM species, chloroplast photosynthesis is not required for the stomatal response to low p CO₂. Stomata did not respond to a decreased p CO₂ in K. daigremontiana in the light when stomata were closed, even when the supply of internal CO₂ was experimentally reduced. We conclude that stomatal closure during phase III is not solely mediated by high internal p CO₂, and suggest that in CAM species the diurnal variability in the responsiveness of stomata to p CO₂ could be explained by hypothesizing the existence of a single CO₂ sensor which interacts with other signalling pathways. When not perturbed by low p CO₂, CO₂ assimilation rate and stomatal conductance were correlated both in the light and in the dark in both species.     

A relationship between carbon dioxide, photosynthetic efficiency and shade tolerance

Net photosynthesis and transpiration of seedlings from shade tolerant, moderately tolerant and intolerant tree species were measured in ambient carbon dioxide (CO₂) concentrations ranging from 312 to 734 ppm. The species used, Fagus grandifolia Ehrh. (tolerant), Quercus alba L., Q. rubra L., Liriodendron tulipifera L. (moderately tolerant), Liquidambar styraciflua L. and Pinus taeda L. (intolerant), are found co-occurring in the mixed pine-hardwood forests of the Piedmont region of the southeastern United States. When seedlings were grown in shaded conditions, photosynthetic CO₂ efficiency was significantly different in all species with the highest efficiency in the most shade tolerant species, Fagus grandifolia , and progressively lower efficiencies in moderately tolerant and intolerant species. Photosynthetic CO₂ efficiency was defined as the rate of increase in net photosynthesis with increase in ambient CO₂ concentration. When plants which had grown in a high light environment were tested, the moderately tolerant and intolerant deciduous species had the highest photosynthetic CO₂ efficiencies but this capacity was reduced when these species grew in low light. The lowest CO₂ efficiency and apparent quantum yield occurred in Pinus taeda in all cases. Water use efficiency was higher for all species in enriched CO₂ environments but transpiration rate and leaf conductance were not affected by CO₂ concentration. High photosynthetic CO₂ efficiency may be advantageous for maintaining a positive carbon balance in the low light environment under a forest canopy.     

Differential responses of mosquito sibling species Anopheles arabiensis and An. quadriannulatusto carbon dioxide, a man or a calf

Field studies on responses of two mosquito sibling species, Anopheles arabiensis Patton and An. quadriannulatus Theobald, to a man, a calf and different release rates of carbon dioxide (man, calf and cow equivalents) were conducted in north-eastern South Africa. Various combinations of baits were compared in two-choice tests, using two mosquito nets, placed 2.5 m apart and 10 cm off the ground. Mosquitoes attracted to the baits were able to enter the nets from below and were collected by means of a suction tube. In a two-choice test between a man and CO₂ (human equivalent, 250 ml/min), 81% of the An. quadriannulatus were caught with CO₂. The reverse was seen for An. arabiensis , where only 20% of the total catch was caught with CO₂ compared to man. High release rates of CO₂ (cow equivalent, 800 ml/min) attracted significantly more An. quadriannulatus than the low release rate (250 ml/min), whereas no significant effect of the release rate of CO₂ on the total catch of An. arabiensis was seen. In the latter species, up to 33% of the attraction of human emanation is attributable to carbon dioxide. Anopheles quadriannulatus was equally attracted to a calf and CO₂ (calf equivalent, 180 ml/min). Catches of other mosquito species showed consistent differences between all treatments which appear to be associated with differences in host-preference, suggesting that the importance of CO₂ in host-seeking behaviour of mosquitoes increases with the degree of zoophily.     

Effect of human odours and positioning of CO2 release point on trap catches of the malaria mosquito Anopheles gambiae sensu stricto in an olfactometer

Abstract.  The anthropophilic malaria mosquito Anopheles gambiae sensu stricto responds to CO₂ and human skin emanations. How these odorants affect the behaviour of this mosquito species is studied in an olfactometer. Carbon dioxide is released either as an homogeneous plume or in a turbulent fashion at two different positions from the trap entrance. Anopheles gambiae is deterred from entering a trap with CO₂ as the only kairomone, when presented as an homogeneous or turbulent plume. This effect is completely overcome by the addition of skin emanations to the CO₂ plume, with a high proportion of mosquitoes found in the trap with skin emanations. Rearrangement of the position of the turbulent CO₂ source so that it is 5 cm downwind of the trap entrance overcomes the deterrent effect of CO₂. Carbon dioxide alone, however, does not elicit higher proportions caught compared with clean air. Further studies with the CO₂ source positioned 5 cm downwind of the trap entrance show that skin emanations alone result in fewer mosquitoes entering the trap than CO₂+ skin emanations. Skin emanations induce more mosquitoes to fly into a trap than a synthetic blend of NH₃+ l-lactic acid when both are combined with CO₂. It is concluded that CO₂ is a poor kairomone when offered alone and that its presence in the plume at the trap entrance deters mosquitoes from entering. By contrast, when positioned just downwind of the trap entrance, CO₂ appears to guide mosquitoes to the vicinity of the trap, where skin emanations then become the principle attractant, causing the mosquito trap entry response. The results of the study have implications for the design of odour-baited traps for this mosquito species.     

Leaf isoprene emission rate as a function of atmospheric CO2 concentration

There is considerable interest in modeling isoprene emissions from terrestrial vegetation, because these emissions exert a principal control over the oxidative capacity of the troposphere. We used a unique field experiment that employs a continuous gradient in CO₂ concentration from 240 to 520 ppmv to demonstrate that isoprene emissions in Eucalyptus globulus were enhanced at the lowest CO₂ concentration, which was similar to the estimated CO₂ concentrations during the last Glacial Maximum, compared with 380 ppmv, the current CO₂ concentration. Leaves of Liquidambar styraciflua did not show an increase in isoprene emission at the lowest CO₂ concentration. However, isoprene emission rates from both species were lower for trees grown at 520 ppmv CO₂ compared with trees grown at 380 ppmv CO₂. When grown in environmentally controlled chambers, trees of Populus deltoides and Populus tremuloides exhibited a 30–40% reduction in isoprene emission rate when grown at 800 ppmv CO₂, compared with 400 ppmv CO₂. P. tremuloides exhibited a 33% reduction when grown at 1200 ppmv CO₂, compared with 600 ppmv CO₂. We used current models of leaf isoprene emission to demonstrate that significant errors occur if the CO₂ inhibition of isoprene is not taken into account. In order to alleviate these errors, we present a new model of isoprene emission that describes its response to changes in atmospheric CO₂ concentration. The model logic is based on assumed competition between cytosolic and chloroplastic processes for pyruvate, one of the principal substrates of isoprene biosynthesis.     

The impact of nitrogen supply on the potential response of a noxious, invasive weed, Canada thistle (Cirsium arvense) to recent increases in atmospheric carbon dioxide

A recognized invasive weed, Canada thistle ( Cirsium arvense L. Scop.) was grown at ambient and pre-ambient concentrations of atmospheric carbon dioxide [CO₂] (373 and 287 μmol mol⁻¹, respectively) at three levels of supplemental nitrogen (N) (3, 6 and 14.5 m M ) from seeding until flowering [77 days after sowing (DAS)]. The primary objective of the study was to determine if N supply limited the potential photosynthetic and growth response of this species to the increase in atmospheric [CO₂] which occurred during the 20th century (i.e. approximately 290 to 370 μmol mol⁻¹ CO₂). Leaf photosynthesis increased both as a function of growth [CO₂] and N supply during the first 46 DAS. Although by 46 DAS photosynthetic acclimation was observed relative to a common measurement CO₂ concentration, there was no interaction with N supply. Both [CO₂] and N increased biomass, relative growth rates and leaf area whereas root : shoot ratio was increased by CO₂ and decreased by increasing N; however, N supply did not effect the relative response to [CO₂] for any measured vegetative parameter up to 77 DAS. Due to the relative stimulation of shoot biomass, total above-ground N increased at elevated [CO₂] for all levels of supplemental N, but nitrogen use efficiency (NUE) did not differ as a function of [CO₂]. Overall, these data suggest that any potential response to increased atmospheric [CO₂] in recent decades for this noxious weedy species was probably not limited by nitrogen supply.     

Adaptation to high CO2 concentration in an optimal environment: radiation capture, canopy quantum yield and carbon use efficiency

The effect of elevated [CO₂] on wheat (Triticum aestivum L. Veery 10) productivity was examined by analysing radiation capture, canopy quantum yield, canopy carbon use efficiency, harvest index and daily C gain. Canopies were grown at either 330 or 1200 μ mol mol^–1[CO₂] in controlled environments, where root and shoot C fluxes were monitored continuously from emergence to harvest. A rapidly circulating hydroponic solution supplied nutrients, water and root zone oxygen. At harvest, dry mass predicted from gas exchange data was 102·8 ± 4·7% of the observed dry mass in six trials. Neither radiation capture efficiency nor carbon use efficiency were affected by elevated [CO₂], but yield increased by 13% due to a sustained increase in canopy quantum yield. CO₂ enrichment increased root mass, tiller number and seed mass. Harvest index and chlorophyll concentration were unchanged, but CO₂ enrichment increased average life cycle net photosynthesis (13%, P < 0·05) and root respiration (24%, P < 0·05). These data indicate that plant communities adapt to CO₂ enrichment through changes in C allocation. Elevated [CO₂] increases sink strength in optimal environments, resulting in sustained increases in photosynthetic capacity, canopy quantum yield and daily C gain throughout the life cycle.     

Modelling the effects of elevated atmospheric CO2 on crown development, light interception and photosynthesis of poplar in open top chambers

An open-top chamber experiment was carried out to examine the likely effects of elevated atmospheric [CO₂] on architectural as well as on physiological characteristics of two poplar clones ( Populus trichocarpa × P. deltoides clone Beaupré and P. deltoides × P. nigra clone Robusta). Crown architectural parameters required as input parameters for a three-dimensional (3D) model of poplar structure, such as branching frequency and position, branch angle, internode length and its distribution pattern, leaf size and orientation, were measured following growth in ambient and elevated [CO₂ ] (ambient + 350 μmol mol^–1) treated open-top chambers. Based on this information, the light interception and photosynthesis of poplar canopies in different [CO₂] treatments were simulated using the 3D poplar tree model and a 3D radiative transfer model at various stages of the growing season. The first year experiments and modelling results showed that the [CO₂] enrichment had effects on light intercepting canopy structure as well as on leaf photosynthesis properties. The elevated [CO₂] treatment resulted in an increase of leaf area, canopy photosynthetic rate and above-ground biomass production of the two poplar clones studied. However, the structural components responded less than the process components to the [CO₂] enrichment. Among the structural components, the increase of LAI contributed the most to the canopy light interception and canopy photosynthesis; the change of other structural aspects as a whole caused by the [CO₂] enrichment had little effect on daily canopy light interception and photosynthesis.     

Origin, fate and significance of CO2 in tree stems

Although some CO₂ released by respiring cells in tree stems diffuses directly to the atmosphere, on a daily basis 15–55% can remain within the tree. High concentrations of CO₂ build up in stems because of barriers to diffusion in the inner bark and xylem. In contrast with atmospheric [CO₂] of c.  0.04%, the [CO₂] in tree stems is often between 3 and 10%, and sometimes exceeds 20%. The [CO₂] in stems varies diurnally and seasonally. Some respired CO₂ remaining in the stem dissolves in xylem sap and is transported toward the leaves. A portion can be fixed by photosynthetic cells in woody tissues, and a portion diffuses out of the stem into the atmosphere remote from the site of origin. It is now evident that measurements of CO₂ efflux to the atmosphere, which have been commonly used to estimate the rate of woody tissue respiration, do not adequately account for the internal fluxes of CO₂. New approaches to quantify both internal and external fluxes of CO₂ have been developed to estimate the rate of woody tissue respiration. A more complete assessment of internal fluxes of CO₂ in stems will improve our understanding of the carbon balance of trees.     

Effects of elevated CO2 concentrations on photosynthesis, growth and reproduction of branches of the tropical canopy tree species, Luehea seemannii Tr. & Planch.

Mature trees have already experienced substantial increases in CO₂ concentrations during their lifetimes, and will experience continuing increases in the future. Small open-top chambers were used to enclose branchlets that were at a height of between 20 and 25 m in the canopy of the tree species Luehea seemannii Tr. & Planch. in a tropical forest in Panamá. Elevated concentrations of CO₂ increased the rate of photosynthetic carbon fixation and decreased stomatal conductance of leaves, but did not influence the growth of leaf area per chamber, the production of flower buds and fruit nor the concentration of nonstructural carbohydrates within leaves. The production of flower buds was highly correlated with the leaf area produced in the second flush of leaves, indicating that the branchlets of mature trees of Luehea seemannii are autonomous to a considerable extent. Elevated levels of CO₂ did increase the concentration of nonstructural carbohydrates in woody stem tissue. Elevated CO₂ concentration also they increased the ratio of leaf area to total biomass of branchlets, and tended to reduce individual fruit weight. These data suggest that the biomass allocation patterns of mature trees may change under future elevated levels of CO₂. Although there were no effects on growth during the experiment, the possibility of increased growth in the season following CO₂ enrichment due to increased carbohydrate concentrations in woody tissue cannot be excluded.     

Leaf Gas Exchange in Canopy Species of a Venezuelan Cloud Forest

Tropical cloud forests are considered humid ecosystems with frequent cloud cover down to the ground surface. However, seasonal variation in precipitation may induce short-term water stress. For canopy leaves, this water stress may also be a consequence of large atmospheric vapor pressure deficits. The objective of this work was to study five canopy cloud forest species to determine if there are restrictions to leaf gas exchange as a consequence of seasonality in precipitation and to daily water deficit due to air evaporative demand mainly during maximum incoming radiation hours. Seasonal daily courses of microclimatic variables (air temperature, relative humidity, photosynthetic photon flux density) and plant responses (leaf water potential, stomatal conductance, CO₂ assimilation rates, leaf nitrogen concentration) were measured at 2400 m asl in Monterrey, an intermontane valley of the Venezuelan Andes. A gradient in terms of responses to water stress conditions was observed between the species, with Clusia multiflora (a 46% reduction in stomatal conductance between seasons) as the most affected and Miconia resimoides (increased stomatal conductance) responding more favorably to slight water stress conditions. If we consider the limitations of water stress and/or light conditions on CO₂ assimilation we may arrange the species into those in which water stress conditions have a greater impact on leaf carbon gain, those where light conditions are determinant and one in which both water stress and light conditions may affect leaf carbon assimilation.     

Role of corticular photosynthesis following defoliation in Eucalyptus globulus

Defoliation can reduce net fixation of atmospheric CO₂ by the canopy, but increase the intensity and duration of photosynthetically active radiation on stems. Stem CO₂ flux and leaf gas exchange in young Eucalyptus globulus seedlings were measured to assess the impact of defoliation on these processes and to determine the potential contribution of re-fixation by photosynthetic inner bark in offsetting the effects of defoliation in a woody species. Pot and field trials examined how artificial defoliation of the canopy affected the photosynthetic characteristics of main stems of young Eucalyptus globulus seedlings. Defoliated potted seedlings were characterized by transient increases in foliar photosynthetic rates and concomitant decreases in stem CO₂ fluxes (both in the dark and light). Defoliated field-grown seedlings showed similar stem CO₂ flux responses, but of reduced magnitude. Despite demonstrating increased re-fixation capability, defoliated potted-seedlings had slowed stem growth. The green stem of seedlings exhibited largely shade-adapted characteristics. Defoliation reduced stem chlorophyll a/b ratio and increased carotenoid concentration. An increased capacity to re-fix internally respired CO₂ (up to 96%) suggested that stem re-fixation represents a previously unexplored mechanism to minimize the impact of foliar loss by maximizing the contribution of all photosynthetic tissues, particularly for young seedlings.