Carnivorous plants are plants that derive some or

Carnivorous plants are plants that derive some or most of their nutrients (but not energy) from trapping and consuming animals or protozoans, typically insects and other arthropods. Carnivorous plants appear adapted to grow in places where the soil is thin or poor in nutrients, especially nitrogen, such as acidic bogs and rock outcroppings. Charles Darwin wrote the first wellknown treatise on carnivorous plants in 1875.

True carnivory is thought to have evolved independently six times in five different orders of flowering plants, and these are now represented by more than a dozen genera. These include about 630 species that attract and trap prey, produce digestive enzymes, and absorb the resulting available nutrients. Additionally, over 300 protocarnivorous plant species in several genera show some but not all these characteristics.

Five basic trapping mechanisms are found in carnivorous plants 1. 2. 3. 4. 5. Pitfall traps (pitcher plants) trap prey in a rolled leaf that contains a pool of digestive enzymes or bacteria. Flypaper traps use a sticky mucilage Snap traps utilize rapid leaf movements Bladder traps suck in prey with a bladder that generates an internal vacuum Lobster-pot traps force prey to move towards a digestive organ with inward-pointing hairs

These traps may be active or passive, depending on whether movement aids the capture of prey. For example, Triphyophyllum is a passive flypaper that secretes mucilage, but whose leaves do not grow or move in response to prey capture. Meanwhile, sundews are active flypaper traps whose leaves undergo rapid acid growth, which is an expansion of individual cells as opposed to cell division. The rapid acid growth allows the sundew tentacles to bend, aiding in the retention and digestion of prey

Pitfall traps Heliamphora, the sun pitche plant. In this genus, the traps are clearly derived evolutionarily from a simple rolled leaf whose margins have sealed together. These plants live in areas of high rainfall in South America such as Mount Roraima and consequently have a problem ensuring their pitchers do not overflow. To counteract this problem, natural selection has favoured the evolution of an overflow similar to that of a bathroom sink—a small gap in the zipped-up leaf margins allows excess water to flow out of the pitcher.

Heliamphora is a member of the Sarraceniaceae, a New World family in the order Ericales(heathers and allies). Heliamphora is limited to South America, but the family contains two other genera, Sarracenia and Darlingtonia, which are endemic to the Southeastern United States (with the exception of one species) and California respectively. S. purpurea subsp. purpurea (the northern pitcher plant) has a more cosmopolitan distribution, found as far north as Canada. Sarracenia is the pitcher plant genus most commonly encountered in cultivation, because it is relatively hardy and easy to grow.

Darlingtonia californica the cobra plant, possesses an adaptation also found in Sarracenia psittacina and, to a lesser extent, in Sarracenia minor: the operculum is balloon-like and almost seals the opening to the tube. This balloon-like chamber is pitted with areolae, chlorophyll-free patches through which light can penetrate. Insects, mostly ants, enter the chamber via the opening underneath the balloon. Once inside, they tire themselves trying to escape from these false exits, until they eventually fall into the tube. Prey access is increased by the "fish tails", outgrowths of the operculum that give the plant its name. Some seedling Sarracenia species also have long, overhanging opercular outgrowths; Darlingtonia may therefore represent an example of neoteny.

The second major group of pitcher plants are the monkey cups or tropical pitcher plants of the genus Nepenthes. In the hundred or so species of this genus, the pitcher is borne at the end of a tendril, which grows as an extension to the midrib of the leaf. Most species catch insects, although the larger ones, particularly N. rajah, also occasionally take small mammals and reptiles. These pitchers represent a convenient source of food to small insectivores. N. bicalcarata possesses two sharp thorns that project from the base of the operculum over the entrance to the pitcher, providing some protection from raids by freeloading mammals.

The pitfall trap has evolved independently in at least two other groups. The Albany pitcher plant Cephalotus follicularis is a small pitcher plant from Western Australia, with moccasin-like pitchers. The rim of its pitcher's opening (the peristome) is particularly pronounced (both secrete nectar) and provides a thorny overhang to the opening, preventing trapped insects from climbing out. The lining of most pitcher plants is covered in a loose coating of waxy flakes, which are slippery for insects, prey that are often attracted by nectar bribes secreted by the peristome and by bright flower-like anthocyanin patterning. In at least one species, Sarracenia flava, the nectar bribe is laced with coniine, a toxic alkaloid also found in hemlock, which probably increases the efficiency of the traps by intoxicating prey.

Flypaper traps The flypaper trap is based on a sticky mucilage, or glue. The leaf of flypaper traps is studded with mucilage-secreting glands, which may be short and nondescript (like those of the butterworts), or long and mobile (like those of many sundews). Flypapers have evolved independently at least five times.

Closely related to Drosera is the Portuguese dewy pine, Drosophyllum, which differs from the sundews in being passive. Its leaves are incapable of rapid movement or growth. Unrelated, but similar in habit, are the Australian rainbow plants (Byblis). Drosophyllum is unusual in that it grows under near-desert conditions; almost all other carnivores are either bog plants or grow in moist tropical areas.

Recent molecular data (particularly the production of plumbagin) indicate that the remaining flypaper, Triphyophyllum peltatum, a member of the Dioncophyllaceae, is closely related to Drosophyllum and forms part of a larger clade of carnivorous and noncarnivorous plants with the Droseraceae, Nepenthaceae, Ancistrocladaceae and Plumbaginaceae. This plant is usually encountered as a liana, but in its juvenile phase, the plant is carnivorous. This may be related to a requirement for specific nutrients for flowering.

The only two active snap traps—the Venus flytrap (Dionaea muscipula) and the waterwheel plant (Aldrovanda vesiculosa)—are believed to have had a common ancestor with similar adaptations. Their trapping mechanism has also been described as a "mouse trap" or "man trap", based on their shape or rapid movement. However, the term snap trap is preferred as other designations are misleading, particularly with respect to the intended prey. Aldrovanda is aquatic and specialised in catching small invertebrates; Dionaea is terrestrial and catches a variety of arthropods, including spiders. Snap traps

The traps are very similar, with leaves whose terminal section is divided into two lobes, hinged along the midrib. Trigger hairs (three on each lobe in Dionaea muscipula, many more in the case of Aldrovanda) inside the trap lobes are sensitive to touch. When a trigger hair is bent, stretch-gated ion channels in the membranes of cells at the base of the trigger hair open, generating an action potential that propagates to cells in the midrib. These cells respond by pumping out ions, which may either cause water to follow by osmosis (collapsing the cells in the midrib) or cause rapid acid growth. The mechanism is still debated, but in any case, changes in the shape of cells in the midrib allow the lobes, held under tension, to snap shut, flipping rapidly from convex to concave and interring the prey. This whole process takes less than a second. In the Venus flytrap, closure in response to raindrops and blown-in debris is prevented by the leaves having a simple memory: for the lobes to shut, two stimul are required, 0. 5 to 30 seconds apart.

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Bladder traps are exclusive to the genus Utricularia, or bladderworts. The bladders pump ions out of their interiors. Water follows by osmosis generating a partial vacuum inside the bladder. The bladder has a small opening, sealed by a hinged door. In aquatic species, the door has a pair of long trigger hairs. Aquatic invertebrates such as Daphnia touch these hairs and deform the door by lever action, releasing the vacuum. The invertebrate is sucked into the bladder, where it is digested. Many species of Utricularia (such as U. sandersonii) are terrestrial, growing on waterlogged soil, and their trapping mechanism is triggered in a slightly different manner. Bladderworts lack roots, but terrestrial species have anchoring stems that resemble them. Temperate aquatic bladderworts generally die back to a resting turion during the winter months, and U. macrorhiza appears to regulate the number of bladders it bears in response to the prevailing nutrient content of its habitat.

Lobster-pot traps A lobster-pot trap is a chamber that is easy to enter, and whose exit is either difficult to find or obstructed by inwardpointing bristles. Lobster pots are the trapping mechanism in Genlisea, the corkscrew plants. These plants appear to specialise in aquatic protozoa. A Y-shaped modified leaf allows prey to enter but not exit. Inward-pointing hairs force the prey to move in a particular direction. Prey entering the spiral entrance that coils around the upper two arms of the Y are forced to move inexorably towards a stomach in the lower arm of the Y, where they are digested. Prey movement is also thought to be encouraged by water movement through the trap, produced in a similar way to the vacuum in bladder traps, and probably evolutionarily related to it.

Borderline carnivores To be a fully fledged carnivore, a plant must attract, kill, and digest prey and it must benefit from absorbing the products of the digestion (mostly amino acids and ammonium ions). To many horticulturalists, these distinctions are a matter of taste. There is a spectrum of carnivory found in plants: from completely non-carnivorous plants like cabbages, to borderline carnivores, to unspecialised and simple traps, like Heliamphora, to extremely specialised and complex traps, like that of the Venus flytrap.

The borderline carnivores include Roridula and Catopsis berteroniana. Catopsis is a borderline carnivorous bromeliad, like Brocchinia reducta. However, unlike B. reducta, which produces the enzyme phosphatase, C. berteroniana has not been shown to produce any digestive enzymes at all. In these pitfall traps, prey simply fall into the urn, assisted by the waxy scales located on the rim. Roridula has a more intricate relationship with its prey.

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