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ISSN : 0974 - 7532 Volume 8 Issue 3 Research & Reviews in BBiiooSScciieenncceess Review RRBS, 8(3), 2014 [94-106] Carnivory in pitcher : An enigmatic meat eating

Niharika R.Patel, R.Krishnamurthy* C.G Bhakta Institute of Biotechnology, Uka Tarsadia University, Mahuva road, Bardoli-394601, Surat, Gujarat, () E-mail : [email protected] ABSTRACT KEYWORDS

Pitcher plants are chiefly carnivorous by nature, as per the name they are families; meat eater plants. They mainly feed on small , small animals and Trapping mechanism; often some small birds which are attracted towards these mysterious meat Digestion; eater plant. They have admirable colors property, sweet secretion, and Importance; production of secondary metabolite for digesting their prey. There habitat Uses. on which they grow in scantily developed and obtain main nutrient through entrapping their prey. They produce extra nitogen and it contains largest group of pitcher plant families. Nepenthaceae and are the best-known and leading groups of pitcher plants and in India three families are known. Involvement of the mechanism and Digestion are related to these two main leading families. For the medicinal purposes they are mostly used and often as ornamental plants they are mostly preferred. Future need of these plants could replace pesticides; these plants could be grown in between the other crops in order to entrap pest and other insects which are mainly responsible for annihilation of crops.  2014 Trade Science Inc. - INDIA

INTRODUCTION tims.[1] Carnivorous plants have adaptations to draw in and Have you ever heard of meat-eating plants? Just trap animals, and absorb nutrients from their bodies. like several animal , various plant species are Most carnivorous plants are moderately diminutive, and — carnivorous that is, they consume insects and other incapable of preying on anything superior than insects “ ” small animals for a primary source of nutrients and min- (the term insectivorous plant is sometimes used), erals for growth. Instead of actually eating insects, car- though a few are known to trap lizards, rodents, and nivorous plants trap them by various means, depending small birds. Carnivorous plants contain chlorophyll and on the kind of plant. After animals such as flies, grass- can use photosynthesis to make carbohydrates, like hoppers, and spiders are trapped, a pool of enzymes common plants. But, they grow in habitats with poor secreted by the plant digests the prey. They are also soil, and benefit from the extra and other nutri- known as pitcher plants, Carnivorous plants, and In- ents that they can obtain from capturing animals. The sectivorous plants. The insectivorous plants often have purpose of carnivory may be to advance the competi- numerous attractions such as dazzling colors, sweet tive advantage of the plant through increased biomass secretion and other prying to lures their innocent vic- or through flowering and set[2-4] RRBS, 8(3) 2014 Niharika R.Patel and R.Krishnamurthy 95 Review Here the lamina is modified into a pitcher like nitrogen and ). Like all carnivorous plants, structure the leaf apex is modified in to the lid of the they grow in habitat where the soil is too poor in miner- pitcher. The colourful closure is lined with a ring of nec- als and/or too acidic for most plants to survive.[11,14] tar glands. The upper region of the pitcher bears many In cultivation, carnivorous plants require special- digestive glands followed by a slippery surface. Its bot- ized conditions. The soil should be acidic and low in tom contains acidic fluid. Insects trapped by the pitcher nutrients: here at the UConn greenhouses, they grow are digested by the enzymes secreted by the glands most species in a mix of five parts peat moss, and absorbed by the pitcher. Carnivorous plants are two parts perlite, and one part horticultural charcoal. found in many parts of the U.S., but are particularly Almost all carnivores do paramount in humid condi- abundant and diverse in some floodplains and small tions, and soil that is constantly wet. The plants should wetlands in the South-east. Pitcher plant available in be watered with rain or deionized water, to avoid min- India is khasiana.[5-10] eral accumulation in the soil. Most species also require Pitcher plants are carnivorous plants whose prey- high intensity light levels, and the bulk of the UConn trapping mechanism features a deep cavity filled with collection is kept in the very sunniest part of the green- liquid known as a pitfall trap. It is far and wide implicit house complex. Carnivorous plants are also grown by pitfall traps evolved by epiascidiation (infolding of the in-vitro propagation method. leaf with the axadial or upper surface becoming the in- Trapping mechanisms of pitcher plants side of the pitcher), with selection pressure favoring more deeply cupped over evolutionary time. The pitcher Carnivorous plants comprise six essential trapping trap evolved in independently in three eudicot lineages mechanisms (among representative genera or species). and one monocot lineage, representing a case of con- Adhesive trap vergent . Some pitcher plant families (such as The simplest trapping mechanism is the adhesive Nepenthaceae) are placed within clades consisting trap. it aiso contains three mechanism Flypaper mostly of flypaper traps, signifying that some pitchers trap(e.g.), Fixed tentacles (e.g., may have evolved from the common ancestors of , , ), and Mobile ’ [11-13] today s flypaper traps by loss of mucilage. tentacles (e.g.) but the flypaper-like leaves of Foraging, flying or crawling insects such as flies are Pinguicula have specific short stalked glands that se- attracted to the cavity formed by the cupped leaf, often crete sticky mucilage that traps miniature creatures. The ’ by visual lures such as anthocyanin pigments, and nec- glands are stalked so the leaf itself doesn t smother in tar bribes. The rim of the pitcher (peristome) is slip- slime while it waits for a prey.[15-21] pery, when moistened by condensation or , caus- ing insects to fall into the entrap. Pitcher plants may Pitfall trap also contain waxy scales, protruding aldehyde crystals, cuticular folds, downward pointing hairs, or guard-cell- originating lunate cells on the inside of the pitcher to ensure that insects cannot climb out. The small bodies of liquid contained within the pitcher traps are called phytotelmata. They drown the , and the body of it is gradually dissolved. This may occur by bacterial ac- tion (the bacteria being washed into the pitcher by rain- fall) or by enzymes secreted by the plant itself. Further- more, some pitcher plants contain mutualistic insect lar- Figure 1 : Pinguicula gigantea leaf showing the residue of vae which feed on trapped prey, and whose excreta fungus gnats. This slippery sticky leaf is an example of an the plant absorbs. Whatever the mechanism of diges- adhesive trap. tion, the prey items are converted into a solution of amino Pitfall trap also have two different mechanism like acids, peptides, phosphates, ammonium and , from Open with pool of water (e.g., Nepen- which the plant obtains its mineral nutrition (particularly thes, Brocchinia, , Catopsis, 96 Carnivory in Pitcher plants: A.n enigmatic meat eating plant RRBS, 8(3) 2014 Review purpurea and S. rosea) and Covered or no pool of water (e.g.Darlingtonia and most Sarracenia). Pitfall traps of pitcher plants leaves are folded into deep, slimy pools filled with digestive enzymes. Pitfall traps are leaves adapted into pit-like structures. Pitfall traps have vastly modified leaves where each leaf have separate trap. The traps may have nectaries, bright colors, or a -like perfume to attract prey. The traps may have hairs to direct prey to the trap opening or cause the prey to fall into the trap. The edge of the trap is usually Figure 3 : leaves have difficult traps. smooth and the inside of the trap waxy. The open pool In the lobster pot part, preys prick the hole in the trap but are traps have pools of water to sink victims. Pitfall traps incapable to find their way out. are thought to have evolved autonomously on at least hairs. This is the way pigeon traps work except thin four occasions. In all-purpose they are phytotelmata, metal rods are used instead of hairs.like e.g.of pigeon water bodies composed or secreted into specialised trap are (Sarracenia psittacina, ).[15-21] containers, and in due course under arrest by plants for assorted functions such as in particular, the trapping and Snap trap digestion of prey[15-21] Lobster pot trap

Figure 4 : Genlisea violacea seedlings showing the fork- shaped Pigeon trap. The white trap is actually a highly modified rolled underground leaf. You need a microscope to “ ” observe the mechanism of the trap. The stomach is up near green leaf-like structures. Figure 2 : Nepenthes hamata leaf. This is an example of a pitfall trap among a pool of water at the bottom. Snap trap (or steel traps) of the and waterwheel plant are hinged leaves that snap shut when Lobster-pot traps of corkscrew plants are twisted activate hairs are touched. Snap traps leave little to the tubular channels lined with hairs and glands. imagination. A prey moves into the trap, brushes against Darlingtonia and Sarracenia psittacina have a trap the trigger hairs, and you see the trap extremely quickly like to the traps used to grab lobsters. A lobster pot has enclose the prey. Unlike hinged snap traps made by an entrance prey can easily find on the exterior and humans, this snap trap bends the halves of the trap to penetrate, but the entrance is tricky to find or exit from surround the prey. The trap closes by almost explo- the inside. like e.g. of lobster trap are (Sarracenia sively intensifying the cells on the outer surface of the psittacina, Darlingtonia)[15-21] leaf. You can confirm this by noting that before the trap Pigeon trap is triggered, each half of the leaf is flat. After being trig- Genlisea plants are established in water or water gered the halves are cupped. The traps will then slowly soaked soil. Primary preys are protozoans. The prey seal and digest the prey, reopening when it is done. [15-21] enter the trap by pushing past inward pointing hairs. (e.g., Dionaea). Once inside the trap they cannot get back out past the Suction trap RRBS, 8(3) 2014 Niharika R.Patel and R.Krishnamurthy 97 Review ing water out of the sealed trap creating what would be considered a vacuum if air were involved. When trig- gered, they work so quickly that the highest speed video cameras show the prey outside the trap in one frame and already sucked inside on the next frame. Once in- doors the prey is digested and the trap rearms itself. (e.g.).[15-21] Evolution and distribution of pitcher plants Carnivorous plants have evolved separately in nu- Figure 5 : Dionaea muscipula leaf is a snap trap. Notice the three trigger hairs on each half of the inner trap surface. merous families. Approximately 600 carnivorous spe- The trap has nectar glands regarding where the red cies are known, belonging to six angiosperm subclasses transitions to green in this trap. and including and eudicotyledon[33,34]. Suction traps distinctive to bladderworts, are enor- These comprise about 600 species that attract and trap mously modified leaves in the shape of a bladder with a prey, produce digestive enzymes, and suck up the re- hinged door lined with trigger hairs. It has been argued sulting available nutrients. Additionally, over 300s the most complex plant leaf on this planet is the Utricu- protocarnivorous plant species in several genera show laria suction trap. The traps ready themselves by pump- some but not all these characteristics. Carnivorous plants occur worldwide, but species richness and abundance – are highest in wet, open, nutrient-poor habitats Guyana Highlands, the southeastern United States, and West- ern [35]. Recently, substantial diversity has been discovered in wet, treeless, ephem- eral-flush vegetation on granitic and gneissic outcrops in parts of West Africa[36,37]. Carnivorous plants have fascinated evolutionary ecologists, botanists, and horti- culturists for centuries. Darwin (1875) provided the first detailed experimental evidence for carnivory in a num- ber of plant genera and established once and for all that true heterotrophy existed in an autotrophic Kingdom.[38] Figure 6 : Utricularia inflata suction trap leaves. The insectivorous plants are found in nine angiosper- TABLE 1 : Digestion mechanism of carnivorous plant[22-32] TABLE 2 : Pitcher plant families and its species[1] Digestion Different Examples Families Species part mechanism Leaf rolls and B. reducta, of tentacles bend for the Neotropics and Brocchinia Drosera increased prey hectioides B. liniflora of Australia and New contact Byblidaceae Leaf surface Leaf rolls or is Guinea and Byblis gigantea one species, Cephalotus follicularis, rolled to contain Pinguicula Cephalotaceae digestion fluid endemic to Australia ’ Drosophyllum, Triphyophyllum peltatum of western Leaf doesn t move Dioncophyllaceae Triphyophyllum Africa Dionaea, about 110 species; Dionaea, Sealed trap - Utricularia Droseraceae Aldrovanda, Drosera,and Pit or Sarracenia, Drosophyllum - stomach Genlisea Lentibulariaceae Genlisea, Utricularia, Pinguicula Nepenthes, Nepenthes, 70 spp. of Southeast Asia Water pool - Nepenthaceae Darlingtonia and Madagascar Commensal Sarracenia, Heliamphora, - Byblis, Roridula Sarraceniaceae organism Darlingtonia 98 Carnivory in Pitcher plants: A.n enigmatic meat eating plant RRBS, 8(3) 2014 Review mous families. But the families Nepenthaceae and plants or monkey cups, is a of carnivorous plants “ Sarraceniaceae are the best-known and largest groups belong to the family Nepenthaceae. The name mon- ” of pitcher plants and in India three families are known. key cups refers to the fact that monkeys have been The families are given in the TABLE 2 observed drinking rainwater from these plants. Nepen- Insectivorous plants of India belong mainly to three thes leaves are differentiated into a photosynthetically families: Droseraceae, Nepenthaceae and active lamina and a pitcher trap and the leaves of these Lentibulariaceae. These families are given below: plants are also mug-shaped organs specialized for at- Nepenthaceae and sarraceniaceae are the best- tracting, capturing, retaining and digesting the prey. The known and largest groups of pitcher plants pitchers frequently consist of different structural and functional zones: lid, peristome, and upper waxy and Nepenthaceae lower glandular zones within the pitcher.[40,41] The Nepenthaceae family contains a single genus, Carnivorous mechanism Nepenthes, containing over 100 species and numerous hybrids and cultivars. In these Old World pitcher plants, Nepenthes Pitcher color, nectar, biochemicals, and TABLE 3 : Indian pitcher plant families, genera, species and its economic uses[5-10] Family Genera Species Economic uses They are important medicinally. Due to the rich content of It contains two organic acids and enzymes. Used in curdling milk. genera D. indica D.pelata is reported to be used by Ayurvedic practitioners 1.Droseraceae 1. Drosera or D. burmanni ‘ ’ in the preparation of gold-bhasma considered an anti- Sundew D. peltata syphilitic and tonic. A yellowish brown crystalline pigment

from D. peltata is used for dyeing silk. 1. A. 2. Aldrovanda - vesiculosa 1. Nepenthes Nepenthes have some ethnomedical uses. Also treat It contains single khasiana cholera. The liquid inside the pitcher is consumed as a 2. Nepenthaceae genera 2. N. remedy for urinary troubles; it is also used as eye drops for 1. treating redness and itching of eyes. About 30 species are known It contains 4 from India U. aurea and genera but 2 are Utricularia U. reticulata are ornamental, especially suitable for aquaria occurring in inflexa and rockeries. U.stellaris is useful 3. Lentibulariaceae India. U. aurea against cough; 1. Utricularia or U. bifida U. caerulae is useful for dressing of wounds; U. bifida is Bladderworts U. caerulae used as a remedy for urinary disease. U. stellaris U. flexuosa U. brachiata 2. Pinguicula 1. P. alpina - the pitchers are borne at the end of tendrils that extend aroma are the primary attractants in pitchers[41-43] Ne- from the midrib of an otherwise unexceptional leaf. So in penthes pitcher resembles an elongated jar with an open the Nepenthaceae, the pitcher arises from the terminal lid, which is moderately filled with fluid that contains segment of the leaf. The plants themselves are often climb- proteases, chitinases, and other enzymes in a watery ers, accessing the canopy of their habitats using the afore- medium.[44-47] This enzyme mix effects the digestion of mentioned tendrils, although others are found on the the trapped prey. These enzymes also play a role in ground in forest clearings, or as epiphytes on tree.[39] inhibiting microbial growth within the pitcher fluid.[44-47] Nepenthes Most Nepenthes are vines or subscandent shrubs in habit, attaching themselves to adjacent vegetation by Nepenthes popularly known as tropical pitcher the use of looped tendrils which build up from the tips RRBS, 8(3) 2014 Niharika R.Patel and R.Krishnamurthy 99 Review of the leaf blades. The pitchers are in turn produced at ground-dwelling herbs whose pitchers begin from a the tips of the tendrils.[48,49] In general, the fluid-filled horizontal . In this family, the entire leaf forms pitchers are produced to trap and digest invertebrate the pitcher. It contains three genera Sarracenia, prey, and Nepenthes are notable for their dimorphism, Heliamphora, and Darlingtonia The species of the i.e., the production of two distinct pitcher types. Young genus Heliamphora, which are popularly known as “ ” “ ” plants produce rosettes of terrestrial or lower pitch- marsh pitchers (or erroneously as sun pitchers), have a ers. These are usually ovoid or globose in form, rest on simple rolled-leaf pitcher, at the tip of which is a spoon- the substrate, and are characterized by the presence of like structure that secretes nectar. They are restricted a pair of vertically-oriented, wing-like processes. It is to areas of high rainfall in South America. The North usually held that these structures serve to channel prey American genus Sarracenia are the trumpet pitchers, from the pitcher base into the mouth, facilitating cap- which have a more complex trap than Heliamphora, ture. However, this has been found not to be the case with an operculum, which prevents excess accumula- with Jack, the only species in tion of rainwater in most of the species. The single spe- which this idea appears to have been tested to date. As cies in the Californian genus Darlingtonia is commonly the plant grows, it may then manufacture a second pitcher known as the cobra plant, due to its possession of an “ ” “ ” “ ” type, the aerial or upper form. These pitchers are in inflated lid with elegant false-exits, and a forked “ ” general funnel-shaped or cylindrical, and lack the wing- tongue , which serves to ferry ants and other prey to like structures. Occasionally, pitchers of intermediate the entrance of the pitcher.[58] form are produced. In some species, the two pitcher types have been shown to target different prey taxa.[50- 56]

Nepenthes khasiana Nepenthes khasiana is a tropical pitcher plant of the genus Nepenthes. It comprises app. 70 species in the world it is the only Nepenthes species native to India and is largely endemic to the Khasi Hills in North East India.[57] and it is used for medicine. Additionally, it is thought to attract prey by emitting ultraviolet light. Figure 7 : Image of Nepenthes khasiana N. khasiana is a short, stout, prostrate to climbing un- der shrub with sub-cylindrical pitchers. These pitchers Sarracenia pitcher plant are modified leaf blades. The basal part of the long peti- Sarracenia is a genus comprising 8 to 11 species ‘ ole is flattened into a leaf blade like structure phyl- of North American pitcher plants, commonly called ’ lode . The distal part of the petiole is a narrow wiry trumpet pitchers. They are mostly grown in southern ’ spring like structure called tendril that coils around sup- United States. The plant s leaves have evolved into a porting plants. The pitchers, colored bright red, green, funnel in order to trap insects, digesting their prey with yellow or a mosaic of all these, dangle in the air. They proteases and other enzymes. The insects are attracted vary greatly in size, ranging from 5 to 30 cm in length. by a nectar-like secretion on the lip of pitchers, as The mouth of the pitcher is furnished with a colorful half well as a combination of color and scent. slimy foot- ’ open lid. Nepenthes confirms to the pitfall type of trap. ing at the pitchers rim, aided in at least one species Nepenthes khasiana is propagated by by a narcotic drug lacing the nectar, causes insects to Micropropagation method by using seed fall inside, where they die and are digested by the plant and nodal explants and a couple of thousand plants have as a nutrient source. Sarracenia are herbaceous pe- been reintroduced in its habitats.[5-10] rennial plants that grow from a subterranean rhizome, Sarraceniaceae with many tubular pitcher-shaped leaves radiating out from the growing point, and then turning upwards with They are New World pitcher plants their trap openings facing the center of the crown. The (Sarraceniaceae), which comprise three genera, are ‘ ’ trap is a vertical tube with a hood (the operculum) 100 Carnivory in Pitcher plants: A.n enigmatic meat eating plant RRBS, 8(3) 2014 Review extending over its entrance; and below it the top of Northern pitcher plant () the tube usually has a rolled lip (the peristome) which They are also well-known as Purple Pitcher Plant. secretes nectar and scents. The hood itself frequently Only S. purpurea usually contains significant amounts produces nectar too, but in lesser quantities. The in- of rainwater in its tubular pitchers. The Purple Pitcher side of the pitcher tube, regardless of species, can be Plant is the most ordinary carnivorous plant in North divided into three to five distinguishable zones: zone 1 America. It can be found in the Florida panhandle all is the operculum (or hood), zone 2 is the peristome the way up to Canada. The plant can vary in color, and rest of the trap entrance, while zones 3 and 4 from lime green to dark purple. Purple Pitcher pro- (which in some species are combined) and 5 (only duces red and purple . Rainwater collects at the present in S. purpurea) are further divisions of the bottom of the tube, drowning any insects that fall in. actual tube. Each of these zones has a specific func- Sarracenia purpurea has a rhizome that produces a – tion, with corresponding morphophysiological char- tuft of 4 10 hollow leaves, forming pitchers that trap [58-62] acteristics. insects and other small prey. The hood is curved around Carnivorous trapping mechanism the mouth of the pitcher. Mechanism of trapping is pit- All Sarracenia trap insects and other prey without fall trap.[1] the use of moving parts. Their traps are static and are based on a combination of lures (including color, scent, and nectar) and inescapability. Most species use a com- bination of scent, drugged nectar, waxy deposits (to clog insect feet) and gravity to topple insect prey into their pitcher. Coniine, an alkaloid drug narcotic to in- sects, has been discovered in the nectar-like secretions of at least S. flava. Once inside, the insect finds the footing very slippery with a waxy surface covering the walls of the pitcher. Further down the tube, downward- Figure 9 : Image of Sarracenia purpurea pointing hairs build retreat impossible, and in the lowest Parrot pitcher plant (S. psittacina) region of the tube, a pool of liquid containing digestive enzymes and wetting agents quickly drowns the prey S. psittacina, the parrot pitcher, uses a lobster-pot and begins digestion. The exoskeletons are usually not style trap that will admit prey (including tadpoles and digested, and over the course of the summer fill up the small fish during floods) but not permit it to find its way pitcher tube.[58-62] out; and sharp inward-pointing hairs force the victim gradually down to the base of the pitcher where it is digested. They are found in wetter areas of low-lying savanna that often flood during heavy rains. The pitch- ers of this species lie flat along the ground and end in hollow, puffed, beaked hoods that are baited with nec- tar. Insects and small animals that enter the tubes are forced to proceed into the pitcher by numerous long needlelike hairs that point toward the base of the leaf, preventing escape.[1] Green hooded pitcher plant () The Green Hooded Pitcher Plant, Sarracenia mi- nor, provides a good example of the pitcher plant way of life. The Green Hooded Pitcher Plant has hollow, Figure 8 : Sarracenia trap insects by pitchers with nectar green, hood-like leaves. The hood prevents the pitcher and slippery footing around the lip. from filling with water. Hooded Pitcher Plants are found RRBS, 8(3) 2014 Niharika R.Patel and R.Krishnamurthy 101 Review ’ between southeastern and the Florida tall. The Yellow Trumpet s nectar has a chemical in it panhandle. They can reach heights between 13 to 19 that produces paralysis when an insect consumes a large inches. Hooded Pitcher Plants produce yellow flowers quantity. Yellow Trumpets () have a that bloom as the first leaves are sprouting. The top of diverse strategy for capturing insects. This pitcher plant “ ” the tube has a canopy or a hood over it. When the does not have hooded leaves. Instead, the vertical hol- insect falls under the shade of the hood, it is alarmed by low leaves will fill with rainwater. Insects attracted to its the darkness and moves deeper into the plant where it nectar will fall in, drown, and be broken down by the is more translucent.[1] mixture of rain water and digestive juices. Yellow Trum- ’ pets flowers have pale yellow, droopy . They grow best in open wet pine savannas.[1]

Figure 10 : Image of Sarracenia psittacina

Sweet pitcher plant () Figure 12 : Image of Sarracenia rubra The Sweet Pitcher Plant (Sarracenia rubra) has White trumpet () narrow, hollow red-veined leaves. This pitcher plant ’ The White Trumpet (Sarracenia leucophylla) is has a fractional hood, which doesn t quite cover the considered by many to be the most stunning of the Ameri- top the leaves, so the pitcher often fills with rainwater. can pitcher plant species. As the name implies, White The flower is a deep red to maroon color on a leafless Trumpets have pitchers that are green toward the bot- stem. It occurs not only in pocosins and wet pine sa- tom while the upper portion of the pitcher and the pitcher vannas but also along Carolina bay margins and in sand lid are bright white, laced with green or red veins. The hill seep areas. Sweet Trumpet grows in northern flowers of this species are large and red. This species Florida; The Sweet Trumpet has minor and weaker has become rare in some areas where it once occurred tubes in the spring. The summer tubes are larger and due to habitat loss and overharvesting for the cut-flower stand tall, from 5 to 19 inches. Sweet Trumpet pro- trade.[1] duces a sweet smell, like a rose. It produces small red flowers in spring.[1]

Figure 11 : Image of Sarracenia minor Figure 13 : Image of Sarracenia flava Yellow trumpet (Sarracenia flava) Green pitcher plant (Sarracenia oreophila) The Yellow Trumpet produces large tall flowers and It is a carnivorous perennial (lives more than 2 years) bright yellow leaves. This plant will grow 20 to 36 inches 102 Carnivory in Pitcher plants: A.n enigmatic meat eating plant RRBS, 8(3) 2014 Review herb with yellowish-green, hollow, pitcher-shaped Guayana Highlands of Venezuela, Guyana, and Brazil, leaves. The pitchers have liquid and enzymes which help Species in the genus Heliamphora are carnivorous the plant digest insects that climb or fall into the plant. plants that consist of a modified leaf form that is fused Short, stiff hairs inside the pitcher pointing downwards into a tubular shape. They have evolved mechanisms to allow insects into the plant but prevent them from crawl- avoid completely filling up with water and attract, trap, ing out. The lid or hood is at the top of the plant. Its and kill insects. At least one species (H. tatei) produces purpose is to stop too much rain water from diluting the its own proteolytic enzymes that allow it to digest its enzymes in the pitcher. The main body is the pitcher prey without the help of symbiotic bacteria. All shaped tube. The plant produces nectar that entices Heliamphora are herbaceous perennial plants that grow insects inside the pitcher. The insects become trapped from a subterranean rhizome. H. tatei grows as a shrub; and are digested by enzymes. The pitcher ranges from up to four meters tall, all other species form prostrate 8-30 inches tall and has purple veins. Basal leaves are rosettes. The leaf size ranges from a few centimeters shorter flat sickle shaped leaves at the bottom of the (H. minor, H. pulchella) up to more than 50 cm (H. plant.[1] ionasi). Heliamphora possess tubular traps formed by rolled leaves with fused edges. Marsh pitcher plants are unusual among pitcher plants in that they lack lids “ ” (opercula), instead having a small nectar spoon on the upper posterior portion of the leaf. This spoon-like structure secretes a nectar-like substance, which serves as a lure for insects and small animals. Each pitcher also exhibits a small slit in its side that allows excess rainwater to drain away, similar to the overflow on a sink. This allows the marsh pitcher plants to maintain a constant maximum level of rainwater within the pitcher. ’ The pitchers inner surface is covered with downward- ’ pointing hairs to force insects into the pitchers lower parts. Heliamphora uses a more typical Figure 14 : Image of Sarracenia leucophylla scheme among flowering plants of having the stigma Heliamphora pitcher plant mature and lose receptivity before the anthers mature ’ The genus Heliamphora contains 18 species of and release pollen. This isn t a detriment to long dis- pitcher plants. The species are collectively known as tance dispersal the way self incompatibility is since a sun pitchers. Scientific name would be marsh pitcher plants. Heliamphora grown nearby savannas in the

Figure 15 : Image of Sarracenia oreophila Figure 16 : Image of Heliamphora chimantensis RRBS, 8(3) 2014 Niharika R.Patel and R.Krishnamurthy 103 Review large plant could have a number of flowers in the ap- the density of organic matter in it, not the relative amount propriate stage to efficiently self pollinate.[63,64] or chemical composition of precipitation. Thus the plant Carnivorous trapping mechanism is not completely passive in its carnivorous metabolism. The vast majority of plants in the genus Heliamphora do not produce their personal digestive enzymes (i.e. proteases, ribonucleases, phosphatases, etc.), relying instead on the enzymes of symbiotic bac- teria to break down their prey. They do, however, at- tract prey through particular visual and chemical signals and trap and kill the prey through a typical pitfall trap. Field studies of H. nutans, H. heterodoxa, H. minor, and H. ionasi have determined that none of these spe- cies produce their own proteolytic enzymes. Most ’ Figure 17 : Image of Heliamphora typically capture ants, while H. tatei s improvements on the design allow it to capture and Importance and uses of pitcher plants [65] absorb nutrients from more flying insects. Pitcher plants are generally used for medicinal pur- Darlingtonia californica poses. Sarracenia purpurea dried-leaf tea used to treat Darlingtonia californica also called the Califor- fever and chills, and ingested to treat smallpox, lung, nia pitcher plant, cobra lily, or cobra plant, is a species and liver ailments. Also used as an aid in childbirth and as of carnivorous plant, the sole member of the genus a diuretic. A study published in 2009 by researchers from Darlingtonia in the family Sarraceniaceae. It is native Tel Aviv University indicates that secretions produced by to Northern California and , growing in carnivorous plants contain compounds (Secondary me- and seeps with cold running water. California pitcher tabolites) that have anti-fungal properties and may lead plant does not grow straight up, but turns left or right on to the development of a new class of anti-fungal drugs its ascent, twisting itself and facing directly opposite from that will be effective against infections that are resistant to current anti-fungal drugs. Pitcher plant is used for diges- its base. Interestingly, each new pitcher grows in such a tive disorders, particularly constipation, urinary tract dis- way so that its final position directs its hood as far away eases, as a diuretic, a cure for smallpox, and to prevent from the others as possible. Not only may this prevent scar formation. By injection, pitcher plant extract (Sarapin) new pitchers from growing into the hoods of older pitch- “ ” is used as a trigger point injection to luxury pain including ers, but it also maximizes the total area surveyed by sciatic pain, intercostal pain, alcoholic or occipital neuri- the hoods, the means by which the plant traps insect tis, brachial plexus neuralgia, meralgia paresthetica, and prey. The hood is the defining feature of the pitcher lumbar or trigeminal neuralgia. Pitcher plant extract plant, in both form and function. The cobra head is the (Sarapin) has been used by injection in mixture with means by which the pitcher plant traps insects and there bupivacaine hydrochloride 0.5% (Marcaine) and gamma by supplements the impoverished soil it grows on. globulin to treat the omohyoideus myofascial pain syn- Darlingtonia is pollinated by a small bee that enters drome. Combinations of injection (extract triamcinolone gaps between the petals and exits (or falls) through the and lidocaine with adrenalin) to treat migraine cephalagia, lower part of flower.[66-68] for diagnosis and dealing of forms of sciatic pain includ- Carnivorous trapping mechanism ing piriformis syndrome (extract and lidocaine), quadra- California pitcher plant does not produce digestive tus lumborum syndrome (extract and corticosteroid), and enzymes to decompose its prey. It relies on the meta- in combination with physiotherapy and an intraoral splint bolic activity of commensal bacteria in its pitcher fluid to treat the Ernest Syndrome that is often mistaken for to break down organic matter in to transportable mol- temporomandibular joint problems. It is also used in pro- ecules, which it can absorb through the pitcher lining. lotherapy (phenol and extract) to cause inflammation at The level of fluid and its acidity seem to be related to the site where the ligaments and tendons attach to the 104 Carnivory in Pitcher plants: A.n enigmatic meat eating plant RRBS, 8(3) 2014 Review bone to stimulate the body to proliferate stronger, shorter, water to create an infusion. This infusion is used in the and less painful ligaments and/or tendons. Pitcher plant treatment of fevers, shakiness, throughout labor and as contains tannins and other chemicals that are thought to a fertility aid in the absence of a menstrual cycle. The assist with some digestive tract problems. There is some infusion was also previously used as a treatment for evidence that suggests that pitcher plant extract may af- smallpox. fect nerves involved in pain sensation. Carnivory pro- Medicinal uses of the vides a means for plants to augment mineral uptake in nutrient-poor environments there by allowing carnivo- Pitcher plant roots can be saturated or steeped in rous plants to compete profitably in such sites. Drocera water to generate a pitcher plant root infusion. Before spp are used in ethno-medicine the bruised leaves, with soaking the roots are cut release to expose their interi- or without ordinary salt, are applied on blisters. Nepen- ors, the place supposed to contain the medicinal com- thes khasiana is use as medicinally. The powder of the pounds. The root infusion was once used to treat small- roots and pitcher is applied in skin diseases, juice from pox. Pitcher plant roots can also be boiled down over unopened pitchers is used as eye-drops against eye dis- a long period of time to produce a decoction. This de- eases and the pitcher with the juice and crushed insects coction is often given to women for the duration of la- and administered to cholera patients. Sarracenia bor to help expel the placenta and after birth. The de- purpurea used as Tonic, laxative, stomachic, diuretic. coction is also used as a treatment for coughing up blood Used in the southern United States in dyspepsia. Native and pulmonary complaints. American Indians used Sarracenia purpurea for a vari- As an ornamental plant ety of medicinal purposes, including as a diuretic and in Pitcher plants in attendance themselves as strange- childbirth, menstruation and treatment of fevers, chills, [5-10] looking and attractive plants. So they are used in gar- whooping cough and smallpox. den as an ornamental plant. Pest control Pitcher plants offer excellent pest control by killing ACKNOWLEDGEMENT bugs such as ants, flies, wasps, bees, beetles, slugs and snails. Some species of pitcher plants can even catch I deeply wish to express my sincere Thanks to John and kill small animals like rats and frogs. Pitcher plants Brittnacher supplying me Images for my review article. “ ” lure bugs into their pitcher, a cylindrical leaf, with nectar I wish to express my gratefulness to DR. R. and scent. The bugs then walk into the slippery cylinder Krishnamurthy for providing me the moral support and trying to reach the nectar, but ultimately slip or become resource for publish my manuscript. The whole contri- tired and fall down into the bottom of the pitcher, which bution for my achievements goes to god, my parents is filled with water, and drown. Once the bug is dead, and my friends who always make my strength and glands located inside the cylinder excrete digestive en- wealth. zymes to liquefy the prey. The pitcher plant then ab- sorbs this nutritive liquid. REFERENCES

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