A simple,light clip‐cage for experiments with aphids

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Does Hypothermic Treatment Provide an Advantage After Spinal Cord Injury Until Surgery? An Experimental Study

We compared the effects of early and late stage hypothermia treatment after spinal cord injury. Five groups each consisting of seven rats were included in this study. In Group 1a (Clip applied-non-treatment group) and Group 1b (Clip applied-treated group) the spinal cords were harvested 1 h after the injury. In Group 2a (clip applied, non-treated group) and Group 2b (clip applied-treated group) the injured segments were harvested 24 h after injury. Group 3 was designed as the sham-operated group. The significantly lower levels of TBARS and GSH-Px in Group 2a, as compared with Group 1b suggests that the hypothermia was effective in the early stage of treatment (P < 0.05). In contrast, TBARS and GSH-Px levels were significantly increased at the 24 h timepoint following treatment (P < 0.05).Short-term systemic hypothermia reduces lipid peroxidation in the early stages after spinal cord injury. This beneficial effect disappears 24 h following systemic hypothermic treatment.     

Plant resistance, plant traits, and host plant choice of the leaf-folding sawfly on the arroyo willow

Abstract. 1. The hypotheses that genetic variation in host plant resistance of the arroyo willow affected leaf folder ( Phyllocolpa sp.) (Hymenoptera: Tenthredinidae) density and that genetic variation in shoot length and leaf length was correlated with resistance were tested.
2. Willows grown in pots and exposed to ovipojsition by the leaf folding sawfly in cages had significantly different densities among clones, indicating variation in resistance caused by genetic differences among conspecific host plants.
3. There was a general correspondence between leaf folder density on potted cuttings and on the plants in the field that were the sources of cuttings.
4. In behavioural choice experiments, susceptible clones (with highest leaf fold densities) had the highest oviposition activity of female leaf folders compared to clones that were resistant to the leaf folder.
5. Clones differed significantly in shoot length and leaf length among clones grown in pots, among clones in the field, and between shoots with galls and shoots without galls on clones in the field.
6. Leaf folder density was significantly positively correlated with mean shoot length on field clones in 1985 and 1986, but was not correlated with leaf length, although leaf length and shoot length were correlated.
7. Leaf length variation among willow clones accounted for a significant portion of the variation in resistance of potted willows, but shoot length was unimportant.     

An experimental study of the effects of predatory fish on macroinvertebrates in a Hong Kong stream

SUMMARY. 1. Predation upon macroinvertebrates by the loach Oreonectes platycephalus Günther (Cobitidae) was studied using predator inclusion/exclusion cages in a series of pools along a Hong Kong stream. Treatments employed were predator exclusion, medium (approximately natural) predator densities (1 fish cage⁻¹) and high predator densities (2 fish cage⁻¹). Macroinvertebrate abundance in cages was monitored after 2 and 4-weeks exposure to predators.
2. The presence of fish was associated with significant declines in the total numbers of macroinvertebrates colonizing cages. However, taxa were influenced differently, with mayflies decreasing by a factor of two while the more mobile shrimps (Atyidae) were unaffected. Chironomid abundance (largely Chironominae) was unaffected by predator density and increased in week 4. Detritus acted as a confounding variable at this time because chironomid abundance was significantly correlated with the weight of accumulated detritus in cages.
3. While invertebrates were more abundant in cages lacking fish, there were no fewer invertebrates in cages with 2 fish than with 1 fish. This may indicate the presence of secure refuges among substrates in the cages, preventing the additional fish from depleting prey further, or a lack of precision of methods due to natural variations in prey densities and spatial patchiness.
4. No significant effects of predators on relative prey abundance or species richness were detected.
5. The impact of predation on prey abundance weakened on week 4, perhaps due to extra refuges among the accumulated detritus. However, drying of the stream increased fish densities in pools so that cages may have become zones of relative safety that were colonized readily by macroinvertebrates. This result highlights the need for year-round investigations to quantify predation effects in Hong Kong's seasonal tropical climate.     

The effect of the water-lily beetle, Galerucella nymphaeae, on leaf production and leaf longevity of the yellow water-lily, Nuphar lutea

SUMMARY. 1. The effect of the water-lily beetle, Galerucella (= Pyrrhalta ) nymphaeae (L.), (Coleoptera, Chrysomelidae), on the leaf production and leaf longevity of the yellow water-lily, Nuphar lutea (= N. luteum ) (L.) Sibth. & Sm., was studied experimentally in southern Finland during one growth season.
2. Twelve cages, each containing a single rosette of the water-lily, were established in the lake during May. Two treatments were randomly applied to cages: (i) control (herbivory not allowed), and (ii) fifty beetles added at the beginning of the season. Leaves were marked individually and the production of new leaves was followed until October.
3. In both treatments the production of floating leaves continued until the end of August but declined towards the end of summer. Herbivory did not affect the number of leaves produced, i.e. the plants did not compensate for the loss of leaves.
4. Leaf longevity was reduced dramatically (up to —60%) when herbivores were present. Leaves that emerged during the most intense period of larval growth had a life-span one-third of that of the control group.     

Stoneflies as ecological engineers – hungry predators reduce fine sediments in stream beds

1. We conducted experiments in a Colorado Rocky Mountain stream to measure the effect of foraging by predatory stoneflies ( Megarcys signata ) on fine sediment deposition and removal.
2. Cages containing one of four treatments were placed in the stream substratum and removed 3 days after fine sediment load to the stream was experimentally increased. Cages containing a stonefly but no prey accumulated less sediment than cages initially with no benthic invertebrates. Although cages with Megarcys plus prey also tended to have less sediment than controls, sediments were not reduced in cages with prey only.
3. Distance from sediment source, depth and current velocity at cages, final prey density and numbers of prey per predator gut at the end of the experiment had no effects on sediment accumulation in cages.
4. To determine the mechanisms underlying this effect, behavioural tests were conducted in a stream-side artificial stream system. To simulate the different hunger levels of stoneflies in cages, Megarcys were fed or starved for 3 days before behavioural trials that were repeated during high (night) and low (day) stonefly feeding periods. During night trials, foraging movements by starved Megarcys were more prolonged and active than those of fed stoneflies, regardless of the presence of prey. However, neither hunger level nor prey presence affected stonefly foraging behaviour during day trials.
5. Results of field experiments suggest that the presence of stoneflies enhances sediment removal from interstitial spaces. Behavioural observations indicate that nocturnal searching for prey by hungry Megarcys dislodges fine sediments from interstitial spaces.
6. Future studies should explore beyond the direct impacts of predators on stream invertebrate communities, and investigate the potential impact of predators on benthic microhabitat quality.     

The solution structure of clip domains from Manduca sexta prophenoloxidase activating proteinase-2

Clip domains are structural modules found in arthropod serine proteinases and some proteolytically inactive homologues, which mediate extracellular signaling pathways of development and immunity. While little is known about their structures or functions, clip domains are proposed to be sites for interactions of proteinases with their activators, cofactors, and substrates. Here we report the solution structure of dual clip domains from Manduca sexta prophenoloxidase activating proteinase-2. Each domain adopts a new mixed alpha/beta fold (a three-stranded antiparallel beta-sheet flanked by two alpha-helices), and the architecture provides structural information on clip domains from a catalytically active proteinase for the first time. Examination of the structure in conjunction with a multiple sequence alignment of the clip domains from different groups suggests a substrate-binding site, a bacteria-interacting region, and a surface for specific interactions. In summary, our results provide insights into the structural basis of clip domain functions and this structure may represent the prototype of group-2 clip domains.     

Stoneflies as ecological engineers – hungry predators reduce fine sediments in stream beds

1. We conducted experiments in a Colorado Rocky Mountain stream to measure the effect of foraging by predatory stoneflies ( Megarcys signata ) on fine sediment deposition and removal.
2. Cages containing one of four treatments were placed in the stream substratum and removed 3 days after fine sediment load to the stream was experimentally increased. Cages containing a stonefly but no prey accumulated less sediment than cages initially with no benthic invertebrates. Although cages with Megarcys plus prey also tended to have less sediment than controls, sediments were not reduced in cages with prey only.
3. Distance from sediment source, depth and current velocity at cages, final prey density and numbers of prey per predator gut at the end of the experiment had no effects on sediment accumulation in cages.
4. To determine the mechanisms underlying this effect, behavioural tests were conducted in a stream-side artificial stream system. To simulate the different hunger levels of stoneflies in cages, Megarcys were fed or starved for 3 days before behavioural trials that were repeated during high (night) and low (day) stonefly feeding periods. During night trials, foraging movements by starved Megarcys were more prolonged and active than those of fed stoneflies, regardless of the presence of prey. However, neither hunger level nor prey presence affected stonefly foraging behaviour during day trials.
5. Results of field experiments suggest that the presence of stoneflies enhances sediment removal from interstitial spaces. Behavioural observations indicate that nocturnal searching for prey by hungry Megarcys dislodges fine sediments from interstitial spaces.
6. Future studies should explore beyond the direct impacts of predators on stream invertebrate communities, and investigate the potential impact of predators on benthic microhabitat quality.     

The effect of microhabitat feeding site selection on aphid foraging and predation risk

Organisms attempt to optimize foraging by maximizing resource acquisition while minimizing predation risk. Aphids (group-living, phloem-feeding insects) routinely change feeding positions and interact with predators and parasites at the single-leaf scale. Here, we assess the life history and predation risk consequences of within-leaf feeding site choices in pea aphids in response to different natural enemies. First, three-chambered clip cages were used to isolate first instar aphids anterior and posterior to a centrally feeding adult on the underside of a single broad bean leaf. Development time to adulthood did not differ between feeding sites, nor did fecundity within the first 24 h of reproduction. Second, we recorded the frequency and latency of natural enemy attacks on aphids adhered to three leaf sites, matching those of the clip cage experiment, on the underside of a single leaf. Aphids feeding nearest the leaf petiole were at greatest risk of predation by a foliar foraging coccinellid predator, Hippodamia convergens, but not by a parasitoid wasp, Aphidius ervi. Thus, feeding nearer the leaf petiole provided no individual life history benefits and exposes the aphid to increased predation risk. We further discuss the notion that feeding at these sites may provide inclusive fitness benefits for colony mates via alarm signaling and subsequent decreased predation success.     

Survival of colour-polymorphic Eristalis arbustorum hoverflies in semi-field conditions

1. The theory on adaptive phenotypic plasticity assumes different fitness optima in different environments. To demonstrate these fitness differences in the field or under laboratory conditions can be difficult.
2. The size of the coloured patches on the abdomen of Eristalis arbustorum varies with rearing temperature. At low temperatures small patches develop and at higher temperatures larger patches develop. Consequently, the colour patch size varies seasonally.
3. To study the relative fitness of two extreme phenotypes in abdominal coloration in the summer, a total of 591 laboratory-reared flies were released in six large population cages. As a measure of fitness, survival was estimated.
4. In all replicate cages the survival of the pale phenotypes was higher than that of the dark phenotypes, indicating a higher fitness for the pale flies under summer conditions.
5. The fat content of flies reared under identical conditions to those released in the cages was measured. Pale flies had ≈ 0·7 mg more fat than dark flies (± 25% extra fat). However, it is argued that this difference in fat reserves could not have been the only cause of the observed difference in survival.
6. The results are discussed in relation to mimicry and thermoregulation.     

Interactive effects of pH and leaf litter on a shredder, the scirtid beetle, Helodes pulchella, inhabiting tree-holes

1. Scirtid beetles (Coleoptera: Scirtidae) are common inhabitants of water-filled treeholes and interact with other detritivores in a processing chain commensalism. The strength of the commensalism is determined by resource quantity and the organisms involved have different tolerances to low pH. To determine the effects and interactions of resource quantity and pH on one of these leaf-shredding scirtid beetles, Helodes pulchella (Guerin), leaf litter and pH were varied independently in experimental microcosms which mimic treehole habitats.
2. Helodes pulchella growth was affected by both resources and pH in two similar experiments conducted in different years. Scirtid larvae grew more slowly in low resource treatments and also grew more slowly in low pH treatments. Scirtid survival was not affected by pH, but was lower in high resource microcosms in the 1997 experiment. Consumption of leaf litter was much higher in high resource microcosms, even though not all leaf litter was consumed in low leaf litter treatments.
3. Thus, water chemistry and leaf litter quantity could have strong effects on resource availability through effects on these beetle facilitators, in turn affecting other insects living in these detritus-based communities.     

Oviposition site selection by Mansonia mosquitoes on water lettuce

Abstract. 1. Viable Mansonia egg masses occur in nature on both the upper and under surfaces of Pistia stratiotes (L.) leaves. Upper-surface masses were from Mansonia dyari Belkin, Heinemann & Page, but lower-surface oviposition was attributable to both M. dyari and M. titillans Walker.
2. Upper-surface egg masses were often out of water but were concentrated near a wet-dry stain line on the leaf. Lower-surface masses were laid underwater within 3 mm of the leaf edge, except for those oviposited through fenestrations caused by insect damage.
3. Oviposition varied seasonally and masses were most numerous between August and December when water lettuce plants were large and crowded. Egg masses were clumped on plants and leaves and concentrated on mature or mature-old leaves that subtended mean angles of 28–34° from the horizontal.
4. On intact plants in cages, M. dyari laid egg masses on both leaf surfaces in approximately the same proportions as observed in nature. On detached leaves floating flat on the water, M. dyari laid all masses on under surfaces.
5. At 20 and 25°C, egg development of M. dyari required 17.4 and 8.8 days (means), these being significantly longer than the mean times for M. titillans. Some larvae from eggs that hatched in the absence of water survived 72 h in the eggshell.
6. Upper-surface oviposition by M. dyari may be an adaptation to the crowded growth of water lettuce, the mosquito's favoured host plant, whose leaves are largely out of water. Leaf ageing and subsidence cause submergence of most upper-surface eggs by the time of hatching.     

The effect of reducing growth in winter wheat on the population dynamics of the grain aphid Sitobion avenae (F.)

1 The effect of reducing the growth of winter wheat on population size and development of the grain aphid Sitobion avenae was studied. 2 Automatic, mobile, crop shading devices, which reduced radiation comparable to levels found on a cloudy day, whilst minimizing other climatic changes, were used to reduce crop growth. Shading was applied between either GS 31 and 39 (1st node detectable → flag leaf ligule just visible) or GS 39 and 55 (flag leaf just visible → 50% of inflorescence emerged). 3 Sitobion avenae populations were initiated at GS 55 and highest populations were subsequently observed on plants shaded between GS 31 and 39. 4 Individual aphids confined within clip cages were observed to measure development from birth to adult moult, adult lifespan and fecundity. Results indicated that adult lifespan was shortest on plants shaded between GS31 and 39, whilst there were no differences in development time or fecundity. 5 Possible explanations for these observations are discussed.     

Decomposition of leaf litter exposed to simulated acid rain in a buffered lotic system

SUMMARY. 1. Exposure to simulated acid rain resulted in changes in the chemical content of riparian vegetation and terrestrial leaf litter and had significant effects on leaf litter decay rates in a well-buffered lotic ecosystem.
2. Foliar nitrogen and phosphorus decreased with decreasing pH of the simulated acid rain and microbial activity was greater on leaf litter exposed to rain of pi I 5.4 than on leaf litter exposed to pH 3.0 or pH 4.0.
3. Detritivore numbers and biomass were significantly higher on leaf litter exposed to pH 5.4, probably due to the greater palatability of the leaf substrate.
4. Decay rates of leaf litter processed in the summer months were significantly lower than decay rates of leaf litter processed during the autumn/winter due to reduction in nitrogen content and microbial respiration.     

Leaf damage effects on leaf expansion timing in Mallotus japonicus (Euphorbiaceae)

For many plant species, biotic factors determining the timing of leaf expansion have not been elucidated sufficiently. We investigated the effects of leaf damage on the timing of leaf defoliation and on the timing of leaf expansion in Mallotus japonicus (Euphorbiaceae). The degree of leaf herbivory of M. japonicus in the field was examined in summer, with subsequent investigation of the date of leaf defoliation in autumn and that of leaf expansion the following spring. Effects of artificial leaf damage on the timing of leaf defoliation and of leaf expansion were also examined in a greenhouse. In the field, the degree of leaf herbivory sustained by M. japonicus did not affect the timing of defoliation. However, the timing of leaf expansion the next spring advanced earlier, concomitantly with the increase of leaf damage during the previous year. Fifty per cent artificial leaf damage in M. japonicus saplings in summer had no marked effect on the timing of leaf defoliation in autumn. However, the effects of leaf damage on the timing of leaf expansion were apparent the following spring: the damaged plants expanded new leaves ca. 8 days earlier than control plants did. Plants that showed earlier leaf expansion tended to have a higher shoot/root ratio. Our results demonstrate that the plants advance the timing of their leaf expansion in response to leaf damage sustained during the previous year, suggesting that the shoot/root ratio is a determining factor.     

Development and testing of a modular strain measurement clip

A novel, multi-use, low-stiffness and low-cost transducer for measuring in vitro strains has been developed and tested. Currently available strain measurement methods are either too expensive, too complicated or too inflexible for multi-use strain measurement. The stainless-steel modular strain measurement clip introduced here was instrumented with four 350 Omega axial strain gauges in a full Wheatstone bridge configuration to take advantage of commonly available strain gauge amplifier equipment. Adjustable extension arms were designed to allow greater application versatility. The clip was calibrated and produced a linear response (R(2)>0.99) over a minimum of 1.04 mm at high amplifier gain. With reduced amplifier settings, testing showed a linear response over a range of 30.5 mm (R(2)>0.99). Clip stiffness was 0.6N/mm of extension arm tip displacement for minimal instrumentation artifact. A validation test was conducted through a comparison of strain clips, surface-mounted strain gauges and theoretical strain in an aluminium rod subjected to axial tensile loading. The two measurement techniques were used to determine the modulus of elasticity of the aluminium rod. Results were within 6% of the known modulus of elasticity for aluminium. A comparative biomechanical test was also performed on an equine third metacarpal specimen. The traditional bonded strain gauging method produced similar results as the new strain clip, but failed to measure ultimate strains since all strain gauges failed prior to specimen failure. Further investigations into the multiple uses of the clip are underway and recommendations for future versions of the clip are given.     

Macroconsumer effects on insect detritivores and detritus processing in a tropical stream

1. Few studies have assessed the effects of macroconsumers, such as fishes and shrimps, on detritus and detritivores.
2. We used an underwater electric field to prevent macroconsumers from feeding in and on leaf packs in a lowland stream in Costa Rica and thus to determine their effects on the density of insect detritivores and decay rates of leaves.
3. Exclusion of macroconsumers resulted in significantly higher densities of small invertebrates inhabiting leaf packs. Most of these were collector–gatherers, none were shredders.
4. Despite the increase in invertebrate density, decay rates of leaves were not statistically different. These findings contrast with results from temperate streams showing that increases in the density of invertebrates in leaf packs typically result in an increased rate of decay.
5. Leaf decay rates and invertebrate densities were also compared between leaf packs placed in electric exclusion treatments and those placed in coarse (2 cm) plastic net bags (as used in many previous studies). Our results suggest that using such netting in tropical streams may deter macroconsumers, which can affect insect density and, potentially, decay rates of organic matter.     

Interactive effects of herbivory and sand burial on growth of a tropical dune plant, Ipomoea pes-caprae

1. This study examined the effects of insect herbivory, sand burial, and the interactive effects of these factors, on the growth of beach morning glory, Ipomoea pes-caprae, a common tropical dune plant. Levels of herbivory and sand burial were manipulated on individual shoots, and effects on stem growth, leaf production, and production of adventitious roots and axillary branches by nodes were examined.
2. Sand burial had a significant positive effect on the production of roots, but did not affect growth in stem length or leaf production.
3. Effects of herbivory were consistently negative, and persisted for 6 weeks after the herbivore damage was incurred. Stem growth rates and leaf production decreased and the production of bare nodes (with no roots or branches) and mortality of apical meristems increased.
4. Interactive effects of herbivory and sand burial influenced both leaf production and root production. In the case of root production, the effects of herbivory in decreasing the proportion of nodes that produced roots occurred only in the presence of burial. In contrast, burial masked the negative effects of herbivory on leaf production.     

Plant leaf area expansion and assimilate production in wheat (Triticum aestivum L.) growing under low phosphorus conditions

Under phosphorus deficiency reductions in plant leaf area have been attributed to both direct effects of P on the individual leaf expansion rate and to a reduced availability of assimilates for leaf growth. In this work we use experimental and simulation techniques to identify and quantify these processes in wheat plants growing under P-deficient conditions. In a glasshouse experiment we studied the effects of soil P addition (0–138 kg P₂O₅ ha^-1) on tillering, leaf emergence, leaf expansion, plant growth, and leaf photosynthesis of wheat plants (cv. INTA Oasis) that were not water stressed. Plants were grown in pots containing a P-deficient (3 mg P g^-1 soil) sandy soil. Sowing and pots were arranged to simulate a crop stand of 173 plants m^-2. Experimental results were integrated in a simulation model to study the relative importance of each process in determining the plant leaf area during vegetative stages of wheat. Phosphorus deficiency significantly reduced plant leaf area and dry weight production. Under P-deficient conditions the phyllochron (PHY) was increased up to a 32%, compared to that of high-P plants. In low-P plants the rate of individual leaf area expansion during the quasi-linear phase of leaf expansion (LER) was significantly reduced. The effect of P deficiency on LER was the main determinant of the final size of the individual leaves. In recently expanded leaves phosphorus deficiency reduced the photosynthesis rate per unit leaf area at high radiation (AMAX), up to 57%. Relative values of AMAX showed an hyperbolic relationship with leaf P% saturating at 0.27%. Relative values of the tillering rate showed an hyperbolic relationship with the shoot P% saturating at values above 0.38%. The value of LER was not related to the concentration of P in leaves or shoots. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the sensitivity of different model variables to changes in model parameters. Simulation results indicated that under mild P stress conditions up to 80% of the observed reduction in plant leaf area was due to the effects of P deficiency on leaf emergence and tillering. Under extreme P-deficient conditions the simulation model failed to explain the experimental results indicating that other factors not taken into account by the model, i.e. direct effects of P on leaf expansion, must have been active. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this work.     

Leaf area expansion and assimilate production in sunflower (Helianthus annuus L.) growing under low phosphorus conditions

Reductions in leaf area and plant growth as a consequence of phosphorus (P) limitations have been attributed both to direct effects of P shortage on leaf expansion rate and to a reduced production of assimilates required for growth. Canopy assimilation and leaf area expansion are closely interrelated processes. In this work we used experimental and simulation techniques to identify and study their importance in determining leaf area on sunflower (Helianthus annuus L.) growing under P-deficient conditions. Experiment 1 was done outdoors, in Buenos Aires, Argentina, and Experiment 2 in a glasshouse in Wageningen, The Netherlands. In both experiments we studied the effects of soil P addition on leaf appearance, leaf expansion, dry matter accumulation, and leaf photosynthesis of non-water stressed plants grown in pots containing a P-deficient soil. Before sowing the equivalent amounts of 0–600 kg of super phosphate ha^-1 were added to the pots. Phosphorus deficiency delayed leaf appearance increasing the value of the phyllochron (PHY) up to 76%, the rate of leaf area expansion during the quasi-linear phase of leaf expansion (LER) was reduced by up to 74%, with respect to high P plants. Phosphorus deficiency reduced by up to 50% the rate of light saturated photosynthesis per unit of leaf area (AMAX) in recently expanded leaves, while at low levels of leaf insertion in the canopy, AMAX was reduced by up to 85%, when compared to that in high P plants. Phosphorus deficiency also reduced the duration of the quasi-linear phase of leaf expansion by up to eight days. The values of LER were related (r = 0.56, P < 0.05) to the mean concentration of P in all the leaves (Leaves P%) and not to the concentration of P in the individual leaf where LER was determined (r = 0.22, P < 0.4) suggesting that under P deficiency individual leaf expansion was not likely to be regulated by the total P concentration at leaf level. The values of AMAX of individual leaves were related (r = 0.79, P < 0.01) to the concentration of total P in the corresponding leaf (Leaf P%). LER showed a hyperbolic relationship with Leaves P% (R² = 0.94, P < 0.01, n = 13) that saturate at 0.14%. AMAX showed a hyperbolic relationship with Leaf P% (R² = 0.73, P < 0.01, n = 53) that saturated with values of Leaf P% higher than 0.22. A morphogenetic model of leaf area development and growth was developed to quantify the effect of assimilate supply at canopy level on total leaf area expansion, and to study the effects of model parameters on the growth of sunflower plants under P-deficient conditions. With this model we identified the existence of direct effects of P deficiency on individual leaf area expansion. However, we calculated that under mild P stress conditions up to 83% of the reduction in the observed leaf area was explained by the particular effects of P% on the rate of leaf appearance, on the duration of the linear period of leaf expansion, and on the value of AMAX. We also calculated that the effects of P deficiency on the value of AMAX alone, explained up to 41% of the observed reductions in total leaf area between the highest and the intermediate P level in Experiment 2. Possible mechanisms of action of the direct effects of P on individual leaf expansion are discussed in this paper.