Effects of light and microcrustacean prey on growth and investment in carnivory in Utricularia vulgaris

SUMMARY 1. In a 5-week enclosure experiment, we studied the effects of light (ambient light, low light) and prey availability (no prey, prey added) on growth and investment in carnivory in Utricularia vulgaris .
2. Investment in carnivory, measured as the proportion of biomass allocated to bladders, was strongly affected by our manipulations of light intensity and prey density. In the treatment with high prey density the light reduction decreased the investment in bladders from 25% to zero. The effect of prey density on investment in bladders was negative. Because prey addition increased the concentration of nutrients, especially phosphorus, we propose that the effect of the prey treatment on investment reflected altered nutrient concentrations.
3. Availability of prey increased growth and apical biomass of Utricularia . As Utricularia had very few bladders in some treatments we suggest that the effect was due to a combination of live prey trapped and increased nutrient availability from dead prey.
4. Abundance of periphyton on Utricularia and on the enclosure walls was highest in the treatments with high prey density where nutrient concentrations were highest. Thus we interpret the response of periphyton as primarily reflecting nutrient availability.     

Nutrient limitation and stoichiometry of carnivorous plants

The cost-benefit model for the evolution of carnivorous plants posits a trade-off between photosynthetic costs associated with carnivorous structures and photosynthetic benefits accrued through additional nutrient acquisition. The model predicts that carnivory is expected to evolve if its marginal benefits exceed its marginal costs. Further, the model predicts that when nutrients are scarce but neither light nor water is limiting, carnivorous plants should have an energetic advantage in competition with non-carnivorous plants. Since the publication of the cost-benefit model over 20 years ago, marginal photosynthetic costs of carnivory have been demonstrated but marginal photosynthetic benefits have not. A review of published data and results of ongoing research show that nitrogen, phosphorus, and potassium often (co-)limit growth of carnivorous plants and that photosynthetic nutrient use efficiency is 20 - 50 % of that of non-carnivorous plants. Assessments of stoichiometric relationships among limiting nutrients, scaling of leaf mass with photosynthesis and nutrient content, and photosynthetic nutrient use efficiency all suggest that carnivorous plants are at an energetic disadvantage relative to non-carnivorous plants in similar habitats. Overall, current data support some of the predictions of the cost-benefit model, fail to support others, and still others remain untested and merit future research. Rather than being an optimal solution to an adaptive problem, botanical carnivory may represent a set of limited responses constrained by both phylogenetic history and environmental stress.     

Effects of zooplankton and conductivity on tropical Utricularia foliosa investment in carnivory

Investment by bladderwort (Utricularia foliosa) in carnivory was estimated in lakes from the Colombian and Brazilian Amazon with different dissolved mineral nutrients and prey availability. As zooplankton abundance in the lake decreased, an increase in the number of bladders per leaf and in the mean bladder size was observed. However, this investment increment in carnivory diminished as the overall availability of dissolved ions in the lake increased. Our results show that carnivorous plants U. foliosa optimise their investment in carnivory, changing bladder number and bladder size according to zooplankton abundance and conductivity.     

Sources of nutrients to rooted submerged macrophytes growing in a nutrient-rich stream

1. The relative contribution of roots and leaves to nutrient uptake by submerged stream macrophytes was tested in experiments where plants were grown in an outdoor flow-channel system. Water was supplied from a nutrient-rich stream with inorganic nitrogen and phosphorus concentrations typical of Danish streams.
2. Four submerged macrophyte species were tested, Elodea canadensis , Callitriche cophocarpa , Ranunculus aquatilis and Potamogeton crispus, and all species were able to satisfy their demand for mineral nutrients by leaf nutrient uptake alone. This was evident from manipulative experiments showing that removal of the roots had no negative impact on the relative growth rate of the plants. Further, the organic N and P concentrations of the plant tissue was constant with time for the de-rooted plants.
3. Enrichment of water and/or sediment had no effect on the relative growth rate of two species, E. canadensis and C. cophocarpa , indicating that in situ nutrient availability was sufficient to cover the needs for growth. Despite the lack of a response in growth rate, a reduced root/shoot biomass ratio was observed with nutrient enrichment of water and/or sediment, and an increased tissue-P concentration in response to open-water enrichment.
4. The open-water nutrient concentrations of the stream in which the experiments were performed are in the upper part of the range found for Danish farmland streams (the majority of Danish streams). Still, however, the negligible effect of nutrient enrichment on the growth of submerged macrophytes observed suggests that mineral nutrient availability might play a minor role in controlling macrophyte growth in most Danish streams.     

Competitive interactions between Nardus stricta L. and Calluna vulgaris (L.) Hull: the effect of fertilizer and defoliation on above- and below-ground performance

1 The role of nutrient supply and defoliation on the competitive interactions between pot-grown Calluna vulgaris and Nardus stricta plants was investigated.WP leading adjustment
2 Young plants were grown alone and together in pots under a combination of fertilizer and defoliation treatments. After 18 months, parameters reflecting both above- and below-ground performance were measured, namely: total above-ground biomass, shoot nitrogen and phosphorus content, root length and the extent of mycorrhizal infection of the roots.
3 In the pots that received fertilizer, the shoot nutrient content and above-ground biomass of Nardus plants increased to a greater extent than those of Calluna plants; this effect was more marked for Nardus plants growing with Calluna plants than for those growing with other Nardus plants. In contrast , Calluna plants growing in competition with Nardus failed to respond to the addition of nutrients. However, in unfertilized pots, Calluna gained more above-ground biomass during the experimental period than Nardus.
4 Calluna had greater root length than Nardus , but Nardus had a higher proportion of its root length infected by mycorrhizal fungi. In both plants, the addition of fertilizer reduced the mycorrhizal infection and increased the root length. Nardus root length was decreased when grown in competition with Calluna only in pots where no nutrients were added. Defoliation decreased the extent of mycorrhizal infection in Calluna roots but not in those of Nardus; defoliation decreased the shoot nutrient content in Calluna plants, but not in Nardus plants.
5 These results suggest that the competitive balance between Nardus and Calluna may be altered by the addition of nutrients, and by defoliation, which may have serious implications for the future dominance of Calluna in heathland ecosystems, particularly those where nutrient inputs are increasing significantly or where grazing pressures are high.     

Seedling growth of four fire-following Mediterranean plant species deprived of single mineral nutrients

1. Anthyllis vulneraria, Cistus creticus, Hippocrepis unisiliquosa and Pinus brutia are frequent post-fire colonizers whose seedlings are subjected to the mineral conditions imposed by ash in the early stages of growth. We test the hypothesis that the effective internal supply of mineral elements in the seeds of these species may complement the external availability of specific nutrients.
2. Newly germinated seedlings were grown in nutrient solutions, each deficient in one of the following elements: nitrogen, phosphorus or potassium. Control treatments consisted of full nutrients and distilled water. Seedling growth was monitored over 12weeks. The final dry masses of the seedlings were taken as a measure of the availability of each element from the seeds' own stored reserves.
3. For the two legume species nitrogen was the most limiting element followed by phosphorus. Potassium deficiency had no effect on the final legume dry biomass. For Cistus, the order was nitrogen followed by potassium, while phosphorus deprivation had no effect on growth during the 12weeks of the experiment. Deprivation of single minerals had no significant effect on seedling growth of Pinus over the same period. The two legumes and Cistus when denied nitrogen had relatively greater relative root ratios than control plants.
4. The relative seedling requirements of each species for early external supplies of the various nutrients for establishment in the field are discussed with reference to the adaptation of the four species to post-fire regeneration.     

The cost of carnivory for Darlingtonia californica (Sarraceniaceae): evidence from relationships among leaf traits

Scaling relationships among photosynthetic rate, foliar nutrient concentration, and leaf mass per unit area (LMA) have been observed for a broad range of plants. Leaf traits of the carnivorous pitcher plant Darlingtonia californica, endemic to southern Oregon and northern California, USA, differ substantially from the predictions of these general scaling relationships; net photosynthetic rates of Darlingtonia are much lower than predicted by general scaling relationships given observed foliar nitrogen (N) and phosphorus (P) concentrations and LMA. At five sites in the center of its range, leaf traits of Darlingtonia were strongly correlated with elevation and differed with soil calcium availability and bedrock type. The mean foliar N : P of 25.2 ± 15.4 of Darlingtonia suggested that these plants were P-limited, although N concentration in the substrate also was extremely low and prey capture was uncommon. Foliar N : P stoichiometry and the observed deviation of Darlingtonia leaf traits from predictions of general scaling relationships permit an initial assessment of the "cost of carnivory" in this species. Carnivory in plants is thought to have evolved in response to N limitation, but for Darlingtonia, carnivory is an evolutionary last resort when both N and P are severely limiting and photosynthesis is greatly reduced.     

The effect of aquatic plant chemistry and morphology on feeding selectivity by the crayfish, Orconectes virilis

SUMMARY. 1. Feeding preference trials conducted with male and female crayfish ( Orconectes virilis ) showed that out of ten submersed aquatic plant species, Chara sp. and Lemna trisulca were the preferred food sources, followed by Myriophyllum exalbescens or Utricularia vulgaris .
2. Analysis of plant fibre, alkaloid, total phosphorus, total and available nitrogen, and organic content showed that crayfish feeding preferences were not related ( P >0.5) to plant fibre or alkaloid content but were negatively correlated ( P <0.05) with plant nutrient content such that species of low nutritional value ( Chara and Lemna ) were consistently preferred.
3. Despite this preference for low nutrient food, daily intake rates by crayfish for phosphorus, nitrogen and organic matter were greatest when feeding on Lemna or Chara due to the large quantity consumed.
4. We suggest that feeding preferences of O. virilis are not determined by plant chemistry but, rather, relate to ease of handling small, short, bottom-dwelling plants such as Chara and Lemna as compared with large erect, roseate or floating-leaved forms. Because crayfish are selective herbivores, their introduction into waters where they do not naturally occur could have a dramatic effect on both the species composition and biomass of submersed plants.     

Relative balance of the cost and benefit associated with carnivory in the tropical Utricularia foliosa

Investment by bladderwort (Utricularia foliosa) in carnivory, in terms of total C and N of bladders per leaf, was estimated in places with different nutrient concentrations from the Yahuarcaca Creek in the Colombian Amazon. The aims were to determine whether nutrient limiting conditions stimulate the investment in carnivory, and the relative balance between C and N invested in carnivory versus C and N obtained from prey. There were no significant differences either for phosphate (PO₄³⁻) concentration or for ammonia (NH₄⁺) concentration among five sampling areas, along approximately 5 km long stretch of the creek, with a pooled mean ± S.D. of 0.19 ± 0.06 and 8.6 ± 3.0 μM, respectively. However, there were significant differences in the nitrate (NO₃⁻) concentration ranging from 0.6 to 2.5 μM. Total C and N of bladders per leaf increased with decreasing NO₃⁻. This corroborates the hypotheses that the carnivorous plant U. foliosa optimises its investment in carnivory according to nutrient availability in the water, and that N is a limiting factor that stimulates the investment in carnivory. The numbers of prey per bladder were also higher under NO₃⁻ limitation, thus enhancing the input of nutrients toward the plant through the bladders. The ratio of total C of prey captured/total C invested in bladders was always lower than 1. However, the efficiency of N was higher since when NO₃⁻ concentration was lower than 1 μM, the ratio of total N of prey captured/total N invested in bladders ranged between 0.97 and 1.67.     

Sediment-related growth limitation of Elodea nuttallii as indicated by a fertilization experiment

1. A fertilization experiment was performed to identify the limiting nutrient for the growth of submerged vegetation in ditches of a peat-grassland system in the Netherlands, in which restoration measures involved ceasing fertilization, exporting nutrients by removal of above-ground plant mass and large-scale introduction of calcium-rich, nutrient-poor artesian water.
2. Growth of Elodea was significantly enhanced by enrichment with nitrogen alone, and by fertilization with nitrogen in combination with phosphorus, and by nitrogen in combination with phosphorus and potassium.
3. Plant tissue nutrient concentrations increased significantly, for nitrogen by enrichment with nitrogen alone, and with nitrogen in combination with phosphorus and potassium; for phosphorus by enrichment with phosphorus alone and with phosphorus in combination with nitrogen and potassium; tissue concentrations of potassium were not enhanced by any treatment.
4. The elemental ratios of treated plants indicated that nitrogen, rather than phosphorus, was limiting in all treatments, except in those involving nitrogen and NK enrichment (when phosphorus was limiting).
5. The efficiency with which plants used nutrients declined with increased supply of nitrogen and phosphorus, but was unchanged when potassium was increased. Efficiencies were similar to those of other aquatic macrophytes.     

Sediment-related growth limitation of Elodea nuttallii as indicated by a fertilization experiment

1. A fertilization experiment was performed to identify the limiting nutrient for the growth of submerged vegetation in ditches of a peat-grassland system in the Netherlands, in which restoration measures involved ceasing fertilization, exporting nutrients by removal of above-ground plant mass and large-scale introduction of calcium-rich, nutrient-poor artesian water.
2. Growth of Elodea was significantly enhanced by enrichment with nitrogen alone, and by fertilization with nitrogen in combination with phosphorus, and by nitrogen in combination with phosphorus and potassium.
3. Plant tissue nutrient concentrations increased significantly, for nitrogen by enrichment with nitrogen alone, and with nitrogen in combination with phosphorus and potassium; for phosphorus by enrichment with phosphorus alone and with phosphorus in combination with nitrogen and potassium; tissue concentrations of potassium were not enhanced by any treatment.
4. The elemental ratios of treated plants indicated that nitrogen, rather than phosphorus, was limiting in all treatments, except in those involving nitrogen and NK enrichment (when phosphorus was limiting).
5. The efficiency with which plants used nutrients declined with increased supply of nitrogen and phosphorus, but was unchanged when potassium was increased. Efficiencies were similar to those of other aquatic macrophytes.     

Nutrient limitation and morphological plasticity of the carnivorous pitcher plant Sarracenia purpurea in contrasting wetland environments

* Plasticity of leaf nutrient content and morphology, and macronutrient limitation were examined in the northern pitcher plant, Sarracenia purpurea subsp. purpurea, in relation to soil nutrient availability in an open, neutral pH fen and a shady, acidic ombrotrophic bog, over 2 yr following reciprocal transplantation of S. purpurea between the wetlands. * In both wetlands, plants were limited by nitrogen (N) but not phosphorus (P) (N content < 2% DW(-1), N : P < 14) but photosynthetic quantum yields were high (F(V)/F(M) > 0.79). Despite carnivory, leaf N content correlated with dissolved N availability to plant roots (leaf N vs , r(2) = 0.344, P < 0.0001); carnivorous N acquisition did not apparently overcome N limitation. * Following transplantation, N content and leaf morphological traits changed in new leaves to become more similar to plants in the new environment, reflecting wetland nutrient availability. Changes in leaf morphology were faster when plants were transplanted from fen to bog than from bog to fen, possibly reflecting a more stressful environment in the bog. * Morphological plasticity observed in response to changes in nutrient supply to the roots in natural habitats complements previous observations of morphological changes with experimental nutrient addition to pitchers.     

Plant morphology and root hydraulics are altered by nutrient deficiency in Pistacia lentiscus (L.)

The plants in arid and semiarid areas are often limited by water and nutrients. Morpho-functional adjustments to improve nutrient capture may have important implications on plant water balance, and on plant capacity to withstand drought. Several studies have shown that N and P deficiencies may decrease plant hydraulic conductance. Surprisingly, studies on the implications of nutrient limitations on water use in xerophytes are scarce. We have evaluated the effects of strong reductions in nitrogen and phosphorus availability on morphological traits and hydraulic conductance in seedlings of a common Mediterranean shrub, Pistacia lentiscus L.. Nitrogen deficiency resulted in a decrease in aboveground biomass accumulation, but it did not affect belowground biomass accumulation or root morphology. Phosphorus-deficient plants showed a decrease in leaf area, but no changes in aboveground biomass. Root length, root surface area, and specific root length were higher in phosphorus-deficient plants than in control plants. Nitrogen and phosphorus deficiency reduced both root hydraulic conductance and root hydraulic conductance scaled by total root surface area. On the other hand, nutrient limitations did not significantly affect root conductance per unit of foliar surface area. Thus, adaptation to low nutrient availability did not affect seedling capacity for maintaining water supply to leaves. The implications for drought resistance and survival during seedling establishment in semi-arid environments are discussed.     

Temporal and spatial change of the investment in carnivory of the tropical Utricularia foliosa

Investment by bladderwort (Utricularia foliosa L.) in carnivory, in terms of biochemical composition (carbohydrates per bladder), elemental composition (carbon and nitrogen per bladder), and morphology of the bladders (length, depth, size of the trap door, and size of antennae), was estimated in seven plants located in Yahuarcaca creek (Colombian Amazon) five times from March to May 2005. The aims were to determine whether investment in carnivory varies temporally (over the growing season of the plant) and/or spatially, and if this potential change in carnivory investment varies according to nutrient conditions. The main differences in the investment in carnivory (changes in bladder number and bladder size, and changes in the size of the antennae) were among locations and there were not important differences over the growing season of the plant. Nitrogen and not phosphorus, was the element that stimulated the investment in carnivory. In addition to changes in bladder number and bladder size, we observed a new strategy to enhance prey capture under nitrogen limitation: changes in the size of the antennae. The size of the antennae was approximately 1.3 higher in those plants located in sites with low NO₃⁻. However, we did not observed changes in the carbon/nitrogen ratio of the bladders or in the relationship between bladder length with bladder depth or size of the trap door. The amount of carbohydrates per bladder was also 1.8 higher in those plants located in sites with low NO₃⁻ (0.13 μM) than those with higher NO₃⁻ concentration (0.39 μM). However, the amount of carbohydrates in the bladder was related with the abundance of periphyton and, hence, it is not possible to conclude that carbohydrate production was a strategy of the plant to enhance the capture of prey. Therefore, our findings do not support the carbohydrate mucilage lure speculations.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

不同林龄刺槐叶片养分重吸收特征及其对土壤养分有效性的响应

为揭示丘陵沟壑区刺槐的养分重吸收特征及其驱动因素,研究该区不同林龄刺槐叶片全氮和全磷的浓度,以及土壤有机碳、全氮、全磷、铵态氮、硝态氮和速效磷浓度及其化学计量,分析了叶片氮磷重吸收效率与土壤养分特性之间的关系。结果表明: 植物和土壤的养分随林龄增长发生显著变化,而土壤总磷和速效磷浓度较低。氮重吸收效率随林分生长先增加后降低,范围为48.2%～54.0%,平均为48.5%;磷重吸收效率则显著增加,范围为45.2%～49.4%,平均为46.9%。氮重吸收效率与土壤氮素和氮磷比呈负向响应,而磷重吸收效率与氮磷比呈显著正相关,与土壤速效磷呈负相关。表明土壤养分有效性的变化负向驱动养分重吸收效率。由于该生境中刺槐林的固氮效应及磷限制,叶片养分重吸收策略对土壤氮磷比响应强烈。     

The response of microbial biomass and hydrolytic enzymes to a decade of nitrogen, phosphorus, and potassium addition in a lowland tropical rain forest

Nutrient availability is widely considered to constrain primary productivity in lowland tropical forests, yet there is little comparable information for the soil microbial biomass. We assessed microbial nutrient limitation by quantifying soil microbial biomass and hydrolytic enzyme activities in a long-term nutrient addition experiment in lowland tropical rain forest in central Panama. Multiple measurements were made over an annual cycle in plots that had received a decade of nitrogen, phosphorus, potassium, and micronutrient addition. Phosphorus addition increased soil microbial carbon (13 %), nitrogen (21 %), and phosphorus (49 %), decreased phosphatase activity by ~65 % and N-acetyl β-glucosaminidase activity by 24 %, but did not affect β-glucosidase activity. In contrast, addition of nitrogen, potassium, or micronutrients did not significantly affect microbial biomass or the activity of any enzyme. Microbial nutrients and hydrolytic enzyme activities all declined markedly in the dry season, with the change in microbial biomass equivalent to or greater than the annual nutrient flux in fine litter fall. Although multiple nutrients limit tree productivity at this site, we conclude that phosphorus limits microbial biomass in this strongly-weathered lowland tropical forest soil. This finding indicates that efforts to include enzymes in biogeochemical models must account for the disproportionate microbial investment in phosphorus acquisition in strongly-weathered soils.     

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

1. The breakdown of oak ( Quercus robur L.), chestnut ( Castanea sativa Miller) and eucalypt ( Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Agüera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days.
2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day^–1 for chestnut leaf litter, from 0.0036 to 0.0051 day^–1 for oak, and from 0.0027 to 0.0158 day^–1 for eucalypt.
3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water.
4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site.
5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.     

Primary nutrients and the biotic integrity of rivers and streams

1. Controls to reduce loadings of primary nutrients to maintain biotic integrity in rivers and streams have not been widely implemented because the relation between nutrients and chlorophyll, and its consequences for higher trophic levels, is confounded in lotic ecosystems by their openness, the variable degree of nutrient limitation and by the effect of physical factors.
2. The relationship between primary nutrients and biotic integrity in rivers and streams was tested using biological, physical and chemical information collected since 1982 from similar locations in streams throughout Ohio using standard procedures.
3. There was a negative correlation between nutrients, especially total phosphorus, and biotic integrity. The deleterious effect of increasing nutrient concentration on fish communities in low order streams was detectable when nutrient concentrations exceeded background conditions (total inorganic nitrogen and phosphorus > 0.61 mg L^–1 and 0.06 mg L^–1, respectively).
4. These results suggest that the control of release of toxins and oxygen‐demanding wastes to rivers is insufficient to protect aquatic life, and confirm the importance of non‐point sources of pollution in catchment planning as well as the combined effect of habitat and riparian quality on nutrient assimilation.     

Effects of supplementary feeding on growth and reproduction of three carnivorous plant species in a subarctic environment

1 The effect of supplementary feeding on growth and reproduction of three carnivorous plants species was investigated over a 6‐year period. Pinguicula alpina , P. villosa and P. vulgaris populations growing at two altitudes in a subarctic environment were fed with fruit flies ( Drosophila melanogaster ).
2 Fed plants increased in size relative to control plants during the first years. Subsequently a stable size difference between feeding levels was established. The weight of the over‐wintering part was higher in fed plants than in control plants.
3 The flowering frequency (i.e. proportion of plants in a population carrying flowers) was also higher in the fed plants. The proportion of flowering plants increased in the feeding treatment compared to control plants during more‐or‐less the whole experimental feeding period (5–6 years). Seed production also increased slowly in response to feeding.
4 No feeding effect on nitrogen and phosphorus concentrations in seeds and over‐wintering parts was found.
5 In most characteristics the high‐altitude populations were less responsive than populations growing at low altitude. In the high‐altitude population reproduction failed repeatedly, suggesting that seed output may be more dependent on abiotic factors than on resource availability in cold environments.
6 Pinguicula villosa and P. vulgaris used the new resources the same summer for increased leaf rosette growth and current reproduction, showing these species to be income breeders, while the third species ( P. alpina ) allocated a larger proportion to storage and future reproduction, characteristic of a capital breeder.