Recent ecophysiological,biochemical and evolutional insights into plant carnivory

BackgroundCarnivorous plants are an ecological group of approx. 810 vascular species which capture and digest animal prey, absorb prey-derived nutrients and utilize them to enhance their growth and development. Extant carnivorous plants have evolved in at least ten independent lineages, and their adaptive traits represent an example of structural and functional convergence. Plant carnivory is a result of complex adaptations to mostly nutrient-poor, wet and sunny habitats when the benefits of carnivory exceed the costs. With a boost in interest and extensive research in recent years, many aspects of these adaptations have been clarified (at least partly), but many remain unknown.ScopeWe provide some of the most recent insights into substantial ecophysiological, biochemical and evolutional particulars of plant carnivory from the functional viewpoint. We focus on those processes and traits in carnivorous plants associated with their ecological characterization, mineral nutrition, cost–benefit relationships, functioning of digestive enzymes and regulation of the hunting cycle in traps. We elucidate mechanisms by which uptake of prey-derived nutrients leads to stimulation of photosynthesis and root nutrient uptake.ConclusionsUtilization of prey-derived mineral (mainly N and P) and organic nutrients is highly beneficial for plants and increases the photosynthetic rate in leaves as a prerequisite for faster plant growth. Whole-genome and tandem gene duplications brought gene material for diversification into carnivorous functions and enabled recruitment of defence-related genes. Possible mechanisms for the evolution of digestive enzymes are summarized, and a comprehensive picture on the biochemistry and regulation of prey decomposition and prey-derived nutrient uptake is provided.     

Effects of competition,litter, and disturbance on an annual carnivorous plant (Utricularia juncea)

The effects of removal of live plants and litter (using herbicide, clipping, and raking) on seedling establishment were examined in Utricularia juncea (bladderwort), an annual carnivorous plant of low frequency in wet, nutrient-poor pine savannas of the southeastern United States. In addition, the spatial distribution of this species in relation to crayfish disturbances was determined. The creation of competition-free gaps in the groundcover canopy in May 1996 (using herbicide) promoted establishment of this species at two sites by September 1997. Standing dead and litter left in herbicide-treated plots inhibited establishment. Density was near zero in undisturbed plots. Natural occurrences of this species were associated with crayfish mounds, which bury plant litter as they erode and increase in area. These results suggest that Utricularia juncea is a fugitive species that depends on disturbances or litter-free microsites to become established in wet, nutrient-poor seepage savannas in southern Mississippi. It is hypothesized that the production of carnivorous traps combined with relatively high allocation to reproductive structures (>90%) and the production of a persistent seed bank make it well-adapted to nutrient-poor and disturbed habitats.     

A novel insight into the cost–benefit model for the evolution of botanical carnivory

Background The cost–benefit model for the evolution of botanical carnivory provides a conceptual framework for interpreting a wide range of comparative and experimental studies on carnivorous plants. This model assumes that the modified leaves called traps represent a significant cost for the plant, and this cost is outweighed by the benefits from increased nutrient uptake from prey, in terms of enhancing the rate of photosynthesis per unit leaf mass or area (A_N) in the microsites inhabited by carnivorous plants.Scope This review summarizes results from the classical interpretation of the cost–benefit model for evolution of botanical carnivory and highlights the costs and benefits of active trapping mechanisms, including water pumping, electrical signalling and accumulation of jasmonates. Novel alternative sequestration strategies (utilization of leaf litter and faeces) in carnivorous plants are also discussed in the context of the cost–benefit model.Conclusions Traps of carnivorous plants have lower A_N than leaves, and the leaves have higher A_N after feeding. Prey digestion, water pumping and electrical signalling represent a significant carbon cost (as an increased rate of respiration, R_D) for carnivorous plants. On the other hand, jasmonate accumulation during the digestive period and reprogramming of gene expression from growth and photosynthesis to prey digestion optimizes enzyme production in comparison with constitutive secretion. This inducibility may have evolved as a cost-saving strategy beneficial for carnivorous plants. The similarities between plant defence mechanisms and botanical carnivory are highlighted.     

Metabolomic analysis reveals reliance on secondary plant metabolites to facilitate carnivory in the Cape sundew,Drosera capensis

Background and AimsSecondary metabolites are integral to multiple key plant processes (growth regulation, pollinator attraction and interactions with conspecifics, competitors and symbionts) yet their role in plant adaptation remains an underexplored area of research. Carnivorous plants use secondary metabolites to acquire nutrients from prey, but the extent of the role of secondary metabolites in plant carnivory is not known. We aimed to determine the extent of the role of secondary metabolites in facilitating carnivory of the Cape sundew, Drosera capensis.MethodsWe conducted metabolomic analysis of 72 plants in a time-series experiment before and after simulated prey capture. We used ultra-high-performance liquid chromatography–tandem mass spectrometry (UHPLC–MS/MS) and the retention time index to identify compounds in the leaf trap tissue that changed up to 72 h following simulated prey capture. We identified associated metabolic pathways, and cross-compared these compounds with metabolites previously known to be involved in carnivorous plants across taxa.Key ResultsFor the first time in a carnivorous plant, we have profiled the whole-leaf metabolome response to prey capture. Reliance on secondary plant metabolites was higher than previously thought – 2383 out of 3257 compounds in fed leaves had statistically significant concentration changes in comparison with unfed controls. Of these, ~34 compounds are also associated with carnivory in other species; 11 are unique to Nepenthales. At least 20 compounds had 10-fold changes in concentration, 12 of which had 30-fold changes and are typically associated with defence or attraction in non-carnivorous plants.ConclusionsSecondary plant metabolites are utilized in plant carnivory to an extent greater than previously thought – we found a whole-metabolome response to prey capture. Plant carnivory, at the metabolic level, likely evolved from at least two distinct functions: attraction and defence. Findings of this study support the hypothesis that secondary metabolites play an important role in plant diversification and adaptation to new environments.     

Energetics and the evolution of carnivorous plants--Darwin's 'most wonderful plants in the world'

Carnivory has evolved independently at least six times in fiveangiosperm orders. In spite of these independent origins, thereis a remarkable morphological convergence of carnivorous planttraps and physiological convergence of mechanisms for digestingand assimilating prey. These convergent traits have made carnivorousplants model systems for addressing questions in plant moleculargenetics, physiology, and evolutionary ecology. New data showthat carnivorous plant genera with morphologically complex trapshave higher relative rates of gene substitutions than do thosewith simple sticky traps. This observation suggests two alternativemechanisms for the evolution and diversification of carnivorousplant lineages. The ‘energetics hypothesis’ positsrapid morphological evolution resulting from a few changes inregulatory genes responsible for meeting the high energeticdemands of active traps. The ‘predictable prey capturehypothesis’ further posits that complex traps yield morepredictable and frequent prey captures. To evaluate these hypotheses,available data on the tempo and mode of carnivorous plant evolutionwere reviewed; patterns of prey capture by carnivorous plantswere analysed; and the energetic costs and benefits of botanicalcarnivory were re-evaluated. Collectively, the data are moresupportive of the energetics hypothesis than the predictableprey capture hypothesis. The energetics hypothesis is consistentwith a phenomenological cost–benefit model for the evolutionof botanical carnivory, and also accounts for data suggestingthat carnivorous plants have leaf construction costs and scalingrelationships among leaf traits that are substantially differentfrom those of non-carnivorous plants. Key words: Carnivorous plants, competition, construction costs, cost–benefit model, Darwin, energetics, niche overlap, phylogeny, prey capture, universal spectrum of leaf traits Received 6 May 2008; Revised 5 June 2008 Accepted 16 June 2008     

Ecophysiological characterization of carnivorous plant roots: Oxygen fluxes,respiration, and water exudation

Various ecophysiological investigations on carnivorous plants in wet soils are presented. Radial oxygen loss from roots of Droseraceae to an anoxic medium was relatively low 0.02 – 0.07 mol(O₂) m^{– 2} s^–1 in the apical zone, while values of about one order of magnitude greater were found in both Sarracenia rubra roots and Genlisea violacea traps. Aerobic respiration rates were in the range of 1.6 – 5.6 mol kg^–1 (f.m.) s^–1 for apical root segments of seven carnivorous plant species and 0.4 – 1.1 mol kg^–1 (f.m.) s^–1 for Genlisea traps. The rate of anaerobic fermentation in roots of two Drosera species was only 5 – 14 % of the aerobic respiration. Neither 0.2 mM NaN₃ nor 0.5 mM KCN influenced respiration rate of roots and traps. In all species, the proportion of cyanide-resistant respiration was high and amounted to 65 – 89 % of the total value. Mean rates of water exudation from excised roots of 12 species ranged between 0.4 – 336 mm 3 kg^–1 (f.m.) s^–1 with the highest values being found in the Droseraceae. Exudation from roots was insensitive to respiration inhibitors. No significant difference was found between exudation rates from roots growing in situ in anoxic soil and those kept in an aerated aquatic medium. Carnivorous plant roots appear to be physiologically very active and well adapted to endure permanent soil anoxia.     

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.     

Root nutrient uptake enhances photosynthetic assimilation in prey-deprived carnivorous pitcher plant <Emphasis Type="Italic">Nepenthes talangensis</Emphasis>

Carnivorous plants grow in nutrient-poor habitats and obtain substantial amount of nitrogen from prey. Specialization toward carnivory may decrease the ability to utilize soil-derived sources of nutrients in some species. However, no such information exists for pitcher plants of the genus Nepenthes, nor the effect of nutrient uptake via the roots on photosynthesis in carnivorous plants is known. The principal aim of present study was to investigate, whether improved soil nutrient status increases photosynthetic efficiency in prey-deprived pitcher plant Nepenthes talangensis. Gas exchange and chlorophyll (Chl) fluorescence were measured simultaneously and were correlated with Chl and nitrogen concentration as well as with stable carbon isotope abundance (δ¹³C) in control and fertilized N. talangensis plants. Net photosynthetic rate (P _N) and maximum- (F_v/F_m) and effective quantum yield of photosystem II (Φ_PSII) were greater in the plants supplied with nutrients. Biomass, leaf nitrogen, and Chl (a+b) also increased in fertilized plants. In contrast, δ¹³C did not differ significantly between treatments indicating that intercellular concentration of CO₂ did not change. We can conclude that increased root nutrient uptake enhanced photosynthetic efficiency in prey-deprived N. talangensis plants. Thus, the roots of Nepenthes plants are functional and can obtain a substantial amount of nitrogen from the soil.     

Hoverflies can sense the risk of being trapped by carnivorous plants: An empirical study using Sphaerophoria menthastri and Drosera toyoakensis

Carnivorous plants are major predators of small insects in some habitats. Because traps of carnivorous plants are serious threats for small insects, it is probable to evolve a mechanism to sense a cue of carnivorous plants and avoid being trapped. However, such a sensing behavior of small insects has never been described. Here we report that a hoverfly species Sphaerophoria menthastri, a major pollinator species of carnivorous sundew Drosera toyoakensis, exhibits a behavior to sense a cue of trap leaves and avoids landing there. In a quadrat (5?m?×?5?m) where D. toyoakensis and other non-carnivorous plant species co-occur, we observed behaviors of hoverflies approaching D. toyoakensis and other plants. The numbers of approaches to trap leaves, flowers of D. toyoakensis, flowers of non-carnivorous Lysimachia fortunei and leaves of Poaceae and Cyperaceae were 9, 60, 52 and 54, respectively, and the numbers of landings to those four organs were 2, 55, 49 and 49, respectively. When S. menthastri approached trap leaves, almost all individuals successfully avoided landing there by 1 or 2 hesitation behaviors. These findings suggest that S. menthastri can sense the cue of trap leaves during an approach.     

Construction costs, payback times, and the leaf economics of carnivorous plants

Understanding how different plant species and functional types "invest" carbon and nutrients is a major goal of plant ecologists. Two measures of such investments are "construction costs" (carbon needed to produce each gram of tissue) and associated "payback times" for photosynthesis to recover construction costs. These measurements integrate among traits used to assess leaf-trait scaling relationships. Carnivorous plants are model systems for examining mechanisms of leaf-trait coordination, but no studies have measured simultaneously construction costs of carnivorous traps and their photosynthetic rates to determine payback times of traps. We measured mass-based construction costs (CC(mass)) and photosynthesis (A(mass)) for traps, leaves, roots, and rhizomes of 15 carnivorous plant species grown under greenhouse conditions. There were highly significant differences among species in CC(mass) for each structure. Mean CC(mass) of carnivorous traps (1.14 ± 0.24 g glucose/g dry mass) was significantly lower than CC(mass) of leaves of 267 noncarnivorous plant species (1.47 ± 0.17), but all carnivorous plants examined had very low A(mass) and thus, long payback times (495-1551 h). Our results provide the first clear estimates of the marginal benefits of botanical carnivory and place carnivorous plants at the "slow and tough" end of the universal spectrum of leaf traits.     

Ecophysiological traits of terrestrial and aquatic carnivorous plants: are the costs and benefits the same?

Identification of tradeoffs among physiological and morphological traits and their use in cost–benefit models and ecological or evolutionary optimization arguments have been hallmarks of ecological analysis for at least 50 years. Carnivorous plants are model systems for studying a wide range of ecophysiological and ecological processes and the application of a cost–benefit model for the evolution of carnivory by plants has provided many novel insights into trait‐based cost–benefit models. Central to the cost–benefit model for the evolution of botanical carnivory is the relationship between nutrients and photosynthesis; of primary interest is how carnivorous plants efficiently obtain scarce nutrients that are supplied primarily in organic form as prey, digest and mineralize them so that they can be readily used, and allocate them to immediate versus future needs. Most carnivorous plants are terrestrial – they are rooted in sandy or peaty wetland soils – and most studies of cost–benefit tradeoffs in carnivorous plants are based on terrestrial carnivorous plants. However approximately 10% of carnivorous plants are unrooted aquatic plants. Here we ask whether the cost–benefit model applies equally well to aquatic carnivorous plants and what general insights into tradeoff models are gained by this comparison. Nutrient limitation is more pronounced in terrestrial carnivorous plants, which also have much lower growth rates and much higher ratios of dark respiration to photosynthetic rates than aquatic carnivorous plants. Phylogenetic constraints on ecophysiological tradeoffs among carnivorous plants remain unexplored. Despite differences in detail, the general cost–benefit framework continues to be of great utility in understanding the evolutionary ecology of carnivorous plants. We provide a research agenda that if implemented would further our understanding of ecophysiological tradeoffs in carnivorous plants and also would provide broader insights into similarities and differences between aquatic and terrestrial plants of all types.     

Peatland establishment on mineral soils: Effects of water level, amendments, and species after two growing seasons

Recent surveys of peatland initiation that occurred over the past 10,000 years in northeastern Alberta have revealed that most peatlands initiated by paludification, or swamping of upland soils. Peatland ecologists have long known the importance of the paludification process, but it has not been transferred to peatland restoration methodologies. We initiated this study to determine if wetland structure and function could be re-established on two well sites established with mineral fill within a peatland complex. At two well sites near Peace River, AB, the mineral material was lowered to near the water level of the surrounding peatland. We placed 288 plots of 2 m × 2 m in size using a series of fertilizer, water level, cultivation, and amendment treatments and then introduced a suite of wetland plants. Four questions are addressed: - (1) Will locally available peatland vascular plant species establish on these wet, compacted, mineral soils? If so: (2) are species responses affected by these treatments? (3) are plants that we did not introduce in the planting regime (weeds) a concern? and (4) will the surrounding bog water chemistry have an effect on water in contact with mineral soils? Results after two growing seasons are - (1) Carex aquatilis and Salix lutea have all successfully established at both well sites; (2) C. aquatilis plants (ramets) have increased to an average of 58.5 per plot, up from the 16 original genets planted; (3) the plant responses to amendments are not significantly different from the control plots; (4) weed abundance is significantly different among some amendment types; and (5) pad ditch water chemistry is affected by the surrounding bog waters.     

Nutrient availability and the carnivorous habit in Utricularia vulgaris

1. Carnivory in plants is thought to enhance growth through an increased supply of nutrients, although there are considerable costs involved. It has been assumed that the relative investment of biomass in traps is inversely proportional to the availability of nutrients from non-carnivorous sources. Our aim was to test the effect of increasing nutrient concentration on investment in carnivory by Utricularia vulgaris .
2. Plants were grown under controlled conditions and nitrogen and phosphorus added at three loadings in a crossed design. Investment in carnivory was assessed as the proportion of (i) leaf biomass and (ii) leaf area comprising traps.
3. There was no effect of nutrient additions on plant growth or periphyton abundance. Investment in carnivory declined with increasing phosphorus loading. There was no effect of nitrogen, despite this being the nutrient commonly thought to be sought by carnivorous plants. Analysis of previously published data also indicated a decline in investment with increasing P availability.
4. Investment in carnivory in U. vulgaris is inversely proportional to the availability of phosphorus from non-carnivorous sources.     

Nepenthes insignis uses a C2-portion of the carbon skeleton of L-alanine acquired via its carnivorous organs, to build up the allelochemical plumbagin

Tropical pitcher plants (Nepenthes) catch animals in their specialized cup-shaped leaves, digest the prey by secreting enzymes, and actively take up the resulting compounds. The benefit of this behaviour is the ability to grow and compete in nutrient-poor habitats. Our present in vitro study shows that not only the nitrogen of alanine fed to the carnivorous organs is used by the plant but that in addition intact C2-units derived from C-2 and C-3 of stable isotope labelled L-alanine serve as building blocks, here exemplarily for the synthesis of the secondary metabolite plumbagin, a potent allelochemical. This result adds a new facet to the benefit of carnivory for plants. The availability of plumbagin by a de novo synthesis probably enhances the plants' fitness in their defence against phytophagous and pathogenic organisms. A missing specific uptake or CoA activation mechanism might be the reason that acetate fed to the pitchers was not incorporated into the naphthoquinone plumbagin. The dihydronaphthoquinone glucosides rossoliside and plumbaside A, here isolated for the first time from Nepenthes, by contrast, showed no incorporation after feeding of any of the two precursors, suggesting these compounds to be storage forms with probably very low turnover rates.     

The Lack of Favorable Responses of an Endangered Pitcher Plant to Habitat Restoration

Restoring habitat structure that existed before active and inadvertent fire suppression is thought to be critical to maintaining populations of some rare plants in fire‐suppressed habitats. Nevertheless, the impacts of habitat restoration on most endangered plants are poorly understood. Current theory predicts and empirical studies have shown that the reduction of shade or competition (frequently a goal of many habitat restoration projects in degraded fire‐dependent ecosystems) benefits plants adapted to nutrient‐poor soils by increasing the benefit‐to‐cost ratio of adaptations for enhanced nutrient capture. Here, I examined how experimental reduction of neighboring plants in a wet longleaf pine community dominated in the ground cover by shrubs and stump sprouts influenced the growth, the reproduction, the carnivorous effort, and the benefits of carnivory in a U.S. federally endangered species, Sarracenia rubra ssp. alabamensis. Two years of data showed no significant effects of neighbor reduction or prey exclusion on any of several indicators of plant performance, nor was there any evidence of a hypothesized morphological trade‐off between shade avoidance and prey capture. These results were unexpected. Inadequate replication and atypical precipitation patterns were ruled out as possible explanations. The population studied here (unlike that of a different, but morphologically similar, species growing in a fire‐maintained pine grass–sedge savanna) did not exhibit the ability to respond to variation in competition from neighboring plants.     

不同生态区域沙地建群种油蒿的钙组分特征

为探索同一物种在不同生态区域钙组分特征的差异,选择我国北方沙地重要建群种油蒿(Artemisia ordosica)为研究对象,采集了内蒙古杭锦旗、乌审旗、阿拉善左旗以及宁夏盐池县和陕西榆林市榆阳区不同沙地类型、不同生长阶段的油蒿样品,利用连续组分法测定分析了油蒿的钙组分特征.结果表明,在油蒿的不同器官中,叶水溶性钙和醋酸溶性钙均显著高于枝和根,叶与根盐酸溶性钙均显著高于枝.在不同生态区域,降水量较多的地区油蒿体内水溶性钙含量较多,降水量较少的地区油蒿体内盐酸溶性钙含量较高.分析得知,降水条件较好的地区较高的水溶性钙主要体现在油蒿的叶中,而降水条件较差的地区较高的盐酸溶性钙主要体现在油蒿的叶和根中.油蒿在不同生长阶段钙组分没有显著差异,但不同类型沙地上油蒿的钙组分却有显著差异.可见,不同生态区域的油蒿,生境条件越好体内水溶性钙含量越高,生境条件越差体内盐酸溶性钙含量越高.     

Photosynthesis and dark respiration of leaves of terrestrial carnivorous plants

Using CO₂ gasometry, net photosynthetic (P _N) and dark respiration rates (R _D) were measured in leaves or traps of 12 terrestrial carnivorous plant species usually grown in the shade. Generally, mean maximum P _N (60 nmol CO₂ g⁻¹(DM) s⁻¹ or 2.7 μmol m⁻² s⁻¹) was low in comparison with that of vascular non-carnivorous plants but was slightly higher than that reported elsewhere for carnivorous plants. After light saturation, the facultatively heliophytic plants behaved as shade-adapted plants. Mean R _D in leaves and traps of all species reached about 50% of maximum P _N and represents the high photosynthetic (metabolic) cost of carnivory.     

Increased nutrient supply facilitates acclimation to high-water level in the marsh plant Deyeuxia angustifolia: The response of root morphology

Both water level and nutrient availability are important factors influencing the growth of wetland plants. Increased nutrient supply might counteract the negative effects of flooding on the growth of the fast-growing species. Experimental evidence is scarce and the mechanism is far from clear. The aim of this study is to identify the role of nutrient availability in acclimation to high-water level by investigating the growth and root morphology of the marsh plant Deyeuxia angustifolia, one of the dominant species in the Sanjiang Plain, China. Experimental treatments included two water levels (0 and 10 cm, relative to soil surface) and three levels of nutrient supply (0, 0.5 and 1 g fertilizer per container). High-water level usually led to decreased biomass accumulation, shoot mass and root mass, whereas biomass accumulation was unaffected by water level at the highest nutrient level, indicating that high-nutrient availability played a role in compensating for the growth loss induced by the high-water level. Increased nutrient supply led to decreased root length in 0 cm water-level treatments, but root length increased with nutrient supply in the 10 cm water-level treatments. High-water level usually led to a lower lateral root density, lateral root:main root length ratio and the diameter of main roots and laterals, whereas increased nutrient supply resulted in thicker main roots or laterals, and a higher total root length, lateral root density and lateral root:main root length ratio. These data indicate that the growth of D. angustifolia is restrained by high-water level, and that increased nutrient supply not only ameliorates root characteristics to acclimate to high-water level but also results in a high-total root length to facilitate nutrient acquisition.     

The Roots of Carnivorous Plants

Carnivorous plants may benefit from animal-derived nutrients to supplement minerals from the soil. Therefore, the role and importance of their roots is a matter of debate. Aquatic carnivorous species lack roots completely, and many hygrophytic and epiphytic carnivorous species only have a weakly devel-oped root system. In xerophytes, however, large, extended and/or deep-reaching roots and sub-soil shoots develop. Roots develop also in carnivorous plants in other habitats that are hostile, due to flood-ing, salinity or heavy metal occurance. Information about the structure and functioning of roots of car- nivorous plants is limited, but this knowledge is essential for a sound understanding of the plants’ physiology and ecology. Here we compile and summarise available information on: (1) The morphology of the roots. (2) The root functions that are taken over by stems and leaves in species without roots or with poorly developed root systems; anchoring and storage occur by specialized chlorophyll-less stems; water and nutrients are taken up by the trap leaves. (3) The contribution of the roots to the nutrient supply of the plants; this varies considerably amongst the few investigated species. We compare nutrient uptake by the roots with the acquisition of nutri-ents via the traps. (4) The ability of the roots of some carnivorous species to tolerate stressful conditions in their habitats; e.g., lack of oxygen, saline conditions, heavy metals in the soil, heat during bushfires, drought, and flooding