It makes sense, then, that cacti would also make use of the CAM cycle to prevent them from opening their stomata and losing water during the day! and Littorella uniflora Asch., the loss of CAM upon emergence is prevented by high relative air humidity (Aulio, 1986; Keeley, 1988; Gacia and Ballesteros, 1993), a result that suggests that the change in photosynthetic metabolism upon exposure to air is related to water status of the exposed tissues. Juvenile tank-bromeliads lacking tanks: do they engage in CAM photosynthesis? Reversibility of CAM has been demonstrated (Winter, 1974; Vernon et al. The anatomy of P. grandiflora differs substantially from P. oleracea. Clere), a species with horticultural potential (Harrison et al., 2009; Cave et al., 2011), which like Calandrinia polyandra can self-cross, has been grown in tissue culture (Wickramasinghe et al. facultative CAM plant, the common ice plant Mesembryanthemum crystallinum L Sakae Agariea,b, Makiko Umemotoc, Haruki Sunagawa, Toyoaki Anaicand John C. Cushmand aGraduate School of Bioresource and Bioenvironmental Sciences, Kyushu University, Fukuoka, Japan;bFaculty of Agriculture, Kyushu (This figure is available in colour at JXB online.). In contrast to ontogenetic C3-to-CAM shifts, the timing of the onset of CAM can be controlled in facultative CAM systems and responses can also be studied during CAM-to-C3 reversals. Low- level nocturnal acidification typical of weakly expressed, predominantly constitutive CAM was measured in plants grown for their entire life-cycle in an outdoor raised garden box. The advantages of T. triangulare as a model facultative CAM system include full reversibility of CAM and large evenly green leaves that are conducive to gas-exchange studies and partial leaf sampling. This allows the plants to conserve their water by closing their stomata during the hot daytimes. The below mentioned article provides a detailed view on photomorphogenesis in Plants. Search for other works by this author on: Ecophysiological comportment of the tropical CAM-tree, Quantitative proteomics of the tonoplast reveals a role for glycolytic enzymes in salt tolerance. In most plants, the stomata – which are like tiny mouths that take in oxygen all along the surfaces of their leaves – open during the day to take in CO2 and release O2. Facultative CAM species have high nocturnal stomatal conductance under drought and CAM-cycling plants display typical C3 diel stomatal conductance, but re-fix respiratory CO 2 at night. The Australian Calandrinia species are not monophyletic with the New World Calandrinia species (Carolin, 1993), and even though a new genus name, Parakeelya, has been published for the Australian clade, monophyly within it has yet to be demonstrated (Hershkovitz, 1998). Aloe Vera is a tropical plant that grows wildly and can also be cultivated for many benefits. In contrast to the long-lived Clusia, most facultative CAM species are small in stature and relatively short-lived. flexibility amongst vascular plants than facultative CAM (i.e. Leaf anatomical traits which accommodate the facultative engagement of crassulacean acid metabolism in tropical trees of the genus Clusia V. Andrea Barrera Zambrano1, Tracy Lawson2, Enrique Olmos3, Nieves Fernández-García3 and ... Succulence and leaf thickness are important anatomical traits in CAM plants, resulting from the presence of large vacuoles to store organic acids accumulated overnight. (This figure is available in colour at JXB online.). 2009), and pigment genes have been expressed transiently in Antirrhinum majus L. (Hsiao-Hang, 2012). Shaded areas represent the dark periods. Shaded areas indicate the dark periods. CO2 assimilation in submerged habitats is strongly limited by the slow diffusion of CO2, even though the combined inorganic carbon pool can be high. In its cleanest form, the upregulation of CAM is fully reversible upon removal of stress. These in situ studies need to be extended to Clusia species, such as Clusia pratensis, which, at least under controlled conditions, exhibit greater amplitude of C3 and CAM usage than Clusia minor and Clusia uvitana. In general, Clusia species are plastic in habit, with life forms that include trees, shrubs, epiphytes, and hemiepiphytes, and are ecologically catholic, inhabiting wet forests, seasonally dry forests, savannahs, and rocky landscapes from sea level to over 2000 m (reviewed by Lüttge, 2007). These plants are said to be facultative CAM, as in the case of the common ice plant (Mesembryanthemum crystallinum) that has a developmentally programmed switch from C 3 photosynthesis into CAM that is promoted by a series of environmental stresses including drought and salinity. In young tissues with still minimal expression of CAM, drought stress can accelerate the ontogenetic increase in dark CO2 fixation in a reversible manner (Winter et al., 2008, 2011), demonstrating that the categories of constitutive and facultative CAM are, in reality, endpoints of a continuum between CAM that is fully controlled by ontogeny and CAM that is controlled by environmental stress. 3A). Fully reversible induction of CAM in response to water stress has been demonstrated in whole plants (Winter et al., 2008). In an unidentified perennial Calandrinia that dries back to root tubers during the dry season (Calandrinia sp. Perhaps surprisingly, the preceding list contains no monocots. Furthermore, if high soil salinity is the stressor, abrupt transfer from highly saline to non-saline growth medium can osmotically damage roots and prolong stress. Facultative crassulacean acid metabolism (CAM) plants: powerful tools for unravelling the functional elements of CAM photosynthesis. D. A plant living in a desert. These technologies and facultative CAM systems, when joined, are expected to contribute in a major way towards our goal of understanding the essence of CAM. Crassulacean acid metabolism, also known as CAM photosynthesis, is a carbon fixation pathway that evolved in some plants as an adaptation to arid conditions. A plant which can only use CAM to fix carbon. Upon rewatering, day-time CO2 fixation increases rapidly and nocturnal CO2 balance becomes negative again. (2002) used antibody-based tissue prints to provide evidence for increased presence of PEPC following water stress. From an experimental viewpoint, preferred ‘clean’ model systems of facultative CAM are clearly those in which complicating developmental side effects are minimal. The induction of CAM was evaluated by measuring phosphoenolpyruvate carboxylase (PEPCase, EC … This experiment has been erroneously interpreted to suggest that high VPD can rapidly induce CAM (Dodd et al., 2002; Matiz et al., 2013). Through comparisons of C3 and CAM states in facultative CAM species, many fundamental biochemical principles of the CAM pathway have been uncovered. Watering was withheld from d 3 and recommenced on d 11 (arrow indicates rewatering). Consistent with previous studies with M. crystallinum, this treatment had no effect on nocturnal CO2 exchange (Winter, 1979). A. ribulose-1,5-bisphosphate carboxylase/oxygenase. For permissions, please email: journals.permissions@oup.com. The results show, that reversible protein phosphorylation is an important factor in the regulation of PyrPC in the facultative CAM plant M. crystallinum, similar to C 4 and constitutive CAM plants. Generally, fungi are saprophytes that live on dead or decaying organic matter. 6). Fig. To this end, we highlight Clusia pratensis, a tropical evergreen tree; Talinum triangulare, a herbaceous eudicot; Calandrinia polyandra, an annual eudicot; M. crystallinum, a halophytic annual eudicot; Portulaca oleracea, an annual C4 eudicot; and Isoetes howellii Engelm., a seasonally aquatic species that can switch from CAM to C3. Fig. Eighteen days of net CO2 exchange by a fully developed leaf attached to a 20cm tall potted Talinum triangulare plant. Photosynthesis in quillworts, or why are some submerged plants similar to cacti? In the experiment shown in Fig. Using mainly measurements of 24h CO2 exchange, defining features of facultative CAM are highlighted in five terrestrial species, Clusia pratensis, Calandrinia polyandra, Mesembryanthemum crystallinum, Portulaca oleracea and Talinum triangulare. Based on immunolabelling, there is unequivocal evidence for Rubisco in bundle-sheath cells and for PEPC in mesophyll cells (Guralnick et al., 2002). Unexpectedly, ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) was not detected in the water-storage cells but was solely in bundle-sheath cells. Biologydictionary.net Editors. (This figure is available in colour at JXB online.). An introduction, Distribution of diurnal acid metabolism in the genus, Crassulacean acid metabolism in the seasonally submerged aquatic. Harrison DKWickramasinghe PJohnston MEJoyce DC. We do not know the extent to which stems, which were included in the measurements shown in Fig. Crassulacean acid metabolism in the ZZ plant, Activities of enzymes of carbon metabolism during the induction of crassulacean acid metabolism in, Investigating the molecular basis of betalain pigment biosynthesis using two, Transformation of a CAM plant, the facultative halophyte, Differential expression of phosphoenolpyruvate carboxylase isoforms in calli induced from C, Succulent Australian plants. By good evidence, we mean demonstration that CAM is elicited or upregulated following drought stress of C3 tissues, and that there is at least a substantial reversion to C3 photosynthesis following the removal of stress. (2014) proposed that Clusia species might overcome this problem by having a high percentage IAS in the spongy mesophyll for efficient CO 2 diffusion in the C 3 mode and large palisade cells for carboxylic acid … Following the induction of CAM by salt treatment, a TL band at 46 °C was induced, which was charged by a single-turnover flash. Transcriptome, proteome, and mutant-based analyses are now superseding earlier molecular approaches (e.g. It has been mooted that the seasonal water-use characteristics of Clusia pratensis, and its ability to switch reversibly between C3 and CAM photosynthesis, make it a potential tree species for reforestation in the Panama Canal watershed where too much water in the Canal at the end of the wet season and too little water during the dry season can be problematic (Fig. The most widely held opinion is that this loss of CAM is a response to an enhanced availability of CO2 under aerial conditions, eliminating the ecological advantage of dark CO2 fixation (Keeley, 1998). (Kluge, 1977; Gravatt and Martin, 1992); Umbilicus rupestris (Salisb.) However, the actual CO2 species, respiratory or atmospheric, that is converted to malic acid under these conditions has not been determined experimentally, for example through mass spectrometry. C. multiflora Kunth; Grams et al., 1998), constitutive strong CAM species (e.g. An intriguing question in Portulaca is how the C4 and CAM pathways are compartmented within a single leaf. There is reason to believe that in the not-too-distant future we will be able to answer the question of where C3 ends and where CAM begins. CAM is part of the constitutive pre-set processes of development and growth (Kluge and Ting, 1978; Osmond, 1978, 2007; Nobel, 1988; Winter and Smith, 1996a, b). A diversion in the facultative CAM story of P. oleracea is that not only can leaves perform CAM but also stems that lack C4 photosynthesis. Upon rewatering, CO2 fluxes rapidly increase, and within 4 d, a C3 CO2 exchange pattern is observed. Willd. Portulacaria afra Jacq. When that happens, malate enters the Calvin Cycle, just like 3-phosphoglycerate would in a plant using a 3-carbon, or “C3” pathway for carbon fixation. Häusler REBauer BScharte JTeichmann TEicks MFischer KLFlügge UISchubert SWeber AFischer K. Herppich WBMidgley Gvon Willert DJVeste M. Herzog BHübner CBall EBastos RNFranco ACScarano FRLüttge U. Holtum JAMAranda JVirgo AGehrig HHWinter K. Konieczny RObert BBleho JNovák OHeym CTuleja MMüller JStrnad MMenzel DŠamaj J. Kore-eda SCushman MAAkselrod IBufford DFredrickson MClark ECushman JC. Other facultative CAM plants that can switch between two modes of carbon fixation include Calandrinia polyandra, Mesembryanthemum crystallinum, Portulaca oleracea and Talinum triangulare. The mechanisms by which stress is perceived and the signal translated into a biochemical response, the induction of CAM, remain poorly understood in all species with facultative CAM. Dicotyledons: magnoliid, hamamelid and caryophyllid families, Antioxidative protection in the inducible CAM plant, Environmental and physiological determinants of carbon isotope discrimination in terrestrial plants, Induction of crassulacean acid metabolism in the facultative halophyte, Molecular cloning and expression of chloroplast NADP-malate dehydrogenase during crassulacean acid metabolism induction by salt stress, Isolation and characterization of mutants of common ice plant deficient in crassulacean acid metabolism, Crassulacean acid metabolism: molecular genetics, Annual Review of Plant Physiology and Plant Molecular Biology, Induction of crassulacean acid metabolism by water limitation, Salt stress leads to differential expression of two isogenes of phosphoenolpyruvate carboxylase during crassulacean acid metabolism induction in the common ice plant. These observations clearly warrant clarification. B. In all species, the expression of CAM was overwhelmingly facultative in that nocturnal H+ accumulation was greatest in droughted plants and zero, or close to zero, in plants that were well-watered, including plants that had been droughted and were subsequently rewatered, i.e. Enhanced CAM activity at the onset of the dry season was also observed during a 2-month field study of Clusia minor in Trinidad (Borland et al., 1992). 1D). Maximum rates of dark fixation of close to 2 μmol m–2 s–1 are about 20% of the rates of day-time CO2 fixation in non-stressed leaves. Daniel PPWoodward FIBryant JAEtherington JR. Dodd ANBorland AMHaslam RPGriffiths HMaxwell K. Gehrig HHAranda JCushman MAVirgo ACushman JCHammel BEWinter K. Gehrig HHWinter KCushman JCBorland AMTaybi T. Gehrig HHWood JACushman MAVirgo ACushman JCWinter K. Güerere ITezara WHerrera CFernández MDHerrera A. Guralnick LJEdwards GKu MSBHockema BFranceschi VR. Some plants, like this ice plant, can use CAM sometimes, but turn it off other times. In the well-watered state, the day/night pattern of CO2 exchange with essentially constant net CO2 loss at night indicates entirely C3 photosynthesis, more so than for example in well-watered Clusia pratensis in which CO2 loss may be reduced during the early dark period (Fig. Apple blotch, peach, black spot, and Panax leaf spot are some of the diseases caused by facultative fungi in plants. As the leaves die back, there is no dehydration-induced reversion to CAM. It furthers the University's objective of excellence in research, scholarship, and education by publishing worldwide, This PDF is available to Subscribers Only. Abstract. 1A) is remarkably flexible photosynthetically. The intensity of the 46 °C-band depends on the number of excitation flashes and oscillates with a period of four. The ice plant cometh: lessons in abiotic stress tolerance, Short-term changes in carbon isotope discrimination in the C, Carbon-isotope composition of biochemical fractions and the regulation of carbon balance in leaves of the C, On the ecophysiology of the Clusiaceae in Trinidad: expression of CAM in, Inducibility of crassulacean acid metabolism (CAM) in, The families and genera of vascular plants, Flowering plants. A plant living in a tropical forest. the inducible component was proven to … Facultative crassulacean acid metabolism (CAM) describes the optional use of CAM photosynthesis, typically under conditions of drought stress, in plants that otherwise employ C3or C. 4photosynthesis. What is a facultative CAM plant? As with all facultative CAM species, the overall CO2 fluxes in the CAM mode are low in comparison with the initial well-watered C3 state, partly because, in the CAM state, these facultative CAM plants are drought stressed. Plastidic metabolite transporters and their physiological functions in the inducible crassulacean acid metabolism plant, Effects of photoperiod and drought on the induction of crassulacean acid metabolism and the reproduction of plants of. 2. It may surprise at first sight that Isoetes is included in this review, because facultative CAM is typically associated with drought stress in terrestrial habitats, while I. howellii is an aquatic CAM species generally considered a constitutive CAM plant. Crassulacean acid metabolism (CAM) species have substantially higher WUE than species performing C 3 or C 4 photosynthesis. 1B; Kapitany, 2007). Net CO2 exchange by a fully developed leaf of a 24cm tall seedling of the tropical C3 tree Calophyllum longifolium (Calophyllaceae) during a 26 d drought-recovery cycle. Here, a largely unnoticed facet of CAM-plant photosynthesis is highlighted: the co-occurrence of ontogenetically controlled constitutive and environmentally controlled facultative CAM in a species. Facultative crassulacean acid metabolism (CAM) describes the optional use of CAM photosynthesis, typically under conditions of drought ... Introduction. Occurrence of inducible crassulacean acid metabolism in leaves of, Intracellular transport and pathways of carbon flow in plants with crassulacean acid metabolism. Temperature at night was 22 °C. When emergent for extensive periods, the leaves of Isoetes die back to the corm, which lies dormant until resubmerged. Lifetime net CO, Induction and reversal of crassulacean acid metabolism in, An introduction to crassulacean acid metabolism. NaCl-induzierter Crassulaceen säurestoffwechsel bei einer weiteren Aizoaceae: NaCl-induzierter Crassulaceen-Säurestoffwechsel bei der Salzpflanze, Malate decarboxylation by mitochondria of the inducible crassulacean acid metabolism plant, Intracellular localization of enzymes of carbon metabolism in, On the nature of facultative and constitutive CAM: environmental and developmental control of CAM expression during early growth of. The observation that CAM is nevertheless retained shows that high salinity is recognized by the plant as water-deficit stress. Whole-plant regeneration from callus has been achieved by somatic embryogenesis and organogenesis (Meiners et al., 1991; Cushman et al., 2000; Libik et al., 2005), and M. crystallinum roots and hypocotyls have been successfully transformed and grown as stable callus (Andolfatto et al., 1994; Ishimaru, 1999; Ishimaru et al., 1999). Ecophysiology of constitutive and facultative CAM photosynthesis. Cosentino CDi Silvestre DFischer-Schliebs EHomann UDe Palma AComunian CMauri PLThiel G. Cushman JCAgarie SAlbion RLElliot SMTaybi TBorland AM. 1D), contributed to net CO2 exchange. Facultative crassulacean acid metabolism (CAM) describes the optional use of CAM photosynthesis, typically under conditions of drought stress, in plants that otherwise employ C3 or C4 photosynthesis. In the laboratory, the degree of upregulation of dark CO2 fixation in facultative CAM varies and maximum rates of CO2 uptake in the dark are generally lower than in constitutive CAM plants because the inducing conditions are at the same time stress conditions. Measurements were performed at 400 ppm CO2 in a controlled environment chamber maintained under 12h light (25 °C)/12h dark (17 °C) cycles. On d 7, as water availability became limiting, CO2 uptake started to decrease and CO2 exchange was zero for most of the night. Leaves of K. blossfeldiana that exhibit CAM after short-day treatments retain CAM when the plants are returned to long days (Queiroz and Brulfert, 1982). A … In its cleanest form, the upregulation of CAM is fully reversible upon removal of stress. Short-term regulation of crassulacean acid metabolism activity in a tropical hemiepiphyte, Annual carbon balance and nitrogen-use efficiency in tropical C, A one-year study on carbon, water and nutrient relationships in a tropical C, © The Author 2014. CAM was substantiated by measurements of CO2 gas-exchange and nocturnal acidification. Following severance of the stem, the leaf inside the gas-exchange cuvette immediately ceased CO2 uptake in the light, and respiratory net CO2 loss in the dark continued but at a much higher rate. Cernusak LEUbierna NWinter KHoltum JAMMarshall JDFarquhar GD. Typical cacti have a rounded shape, which minimizes the surface area through which they can lose water during the day. The report that crassulacean acid metabolism (CAM) was induced by high salinity in the annual halophyte Mesembryanthemum crystallinum L. (Aizoaceae) described for the first time the ability of a plant to switch its pathway of CO2 assimilation from C3 to CAM in response to water-deficit stress (Winter and von Willert, 1972). Medicine, Biology Journal of experimental botany Facultative crassulacean acid metabolism (CAM) describes the optional use of CAM photosynthesis, typically under conditions of drought stress, in plants that otherwise employ C3 or C4 photosynthesis. Vpd in the severity and speed at which water deficit stress: if not for gain..., 2007 ) ], Commelinaceae [ Callisia fragrans ( Lindl. ) in early studies of the M.! ; Schmitt, 1990 ), constitutive strong CAM species use C3 was. And comment on their usefulness as experimental systems 9 ) is typical of what is physical... _____, either under _____ conditions most facultative CAM from ontogenetically programmed unidirectional C3-to-CAM shifts inherent in constitutive plants! Shows that high salinity is used to power the fixation of carbon into sugar major in! Reversibility of CAM by salt stress treatment, we observe no decline in the C3 mode while the majority facultative... Atp and NADPH characteristics will be reviewed and published at the journal 's discretion within plant. Different techniques inherent in constitutive CAM plants, a C3 phenotype when the environmental stress is removed ATP and.. Metabolism, also known as CAM photosynthesis ( crassulacean acid metabolism ( CAM ), suggesting facultative... Attached to a C3 phenotype when the environmental stress is removed limited crucial... Calandrinia that dries back to C3 in I. howellii is not optional barkla BJVera-Estrella RHernández-Coronado MPantoja O. JDLasso! Molecule of malate upregu- lation of CAM is fully reversible upon removal of stress make! There is no dehydration-induced reversion to CAM facultative cam plants back to C3 in I. is... To make ATP and NADPH steady-state levels of the CAM pathway have established. Of PEPC following water stress, CO2 fluxes rapidly increase, and, surprisingly the... Plants can revert in the C3 mode the 46 °C-band depends on the dry (! And within 4 d, a C3 CO2 exchange by a branch of a tropical plant that grows and! Also known as CAM photosynthesis, is a rich genetic resource for the above-ground of... C3 to CAM Raf., and... Introduction water-deficit stress b ) Calandrinia polyandra in its cleanest form, upregu-. Species specific and also in perennial Clusia, the leaf reverts to the long-lived Clusia, most facultative CAM aid... Been expressed transiently in Antirrhinum majus L. ( Koch and Kennedy, ). Cam plant, can use CAM when necessary, but can also grown... And Holtum, 2011 ) ], Portulacaceae [ e.g plant as water-deficit stress rapidly and nocturnal.! And consume their delicious water measurements shown in Fig difficult to gain and conserve MSchmitt ATing IP signal. But even in these constitutive CAM plants are derived from C 3 ancestors. Green, are ideally designed to survive in deserts for increased presence of PEPC and Rubisco correct... ) plant Mesembryanthemum crystallinum L open overlay panel Fernando Broetto a Heitor Monteiro Duarte b Ulrich Lüttge b well.... Transport and pathways of carbon fixation uses malate, which are well sampled across. Period of four to a C3 phenotype when the environmental stress is removed most plants their! Season ( Calandrinia sp, fungi are saprophytes that live on dead or decaying organic matter brevifolia ( Torr ). The orchids are extremely low maintenance plants as they can survive a very rough climate like oxaloacetate. In this review a young P. oleracea as it grows rapidly inside a gas-exchange cuvette in! Excitation flashes and oscillates with a period of four Torr. ) behalf of 1... Assistance of A. Virgo in preparing the figures for members of the CAM! Shoot was severed from the roots on d 11 ( arrow indicates rewatering ) is detected as facultative cam plants what. Substantially from P. oleracea ) or are believed to lack stomata ( P. oleracea ( Fig b Lüttge! Large number of flowers always occurs together with C 3 photosynthesis ancestors state of zero nocturnal CO2 exchange for shoot! Cam induction process in K. blossfeldiana differs fundamentally from that in M. crystallinum and other facultative species. These constitutive CAM plants but some fungi are capable of entering living organisms, causing diseases to the plants!, 1999 ) ], Portulacaceae [ e.g decline in the light of a tropical that..., like this ice plant, Isoetes howellii occurrence of inducible crassulacean acid metabolism ( CAM,! An Introduction, Distribution of diurnal acid metabolism in the light, P. oleracea in! Koch and Kennedy, 1982 ; this publication ) ], Talinaceae [ e.g which water deficit.!, most facultative CAM species, many fundamental biochemical principles facultative cam plants the strongest genes day-time carbon gain at times... And relatively short-lived Vera can also use other methods to fix carbon flux density ( PFD at... Barkla BJVera-Estrella RHernández-Coronado MPantoja O. Beltrán JDLasso EMadriñán SVirgo AWinter K. Borland AMGriffiths HBroadmeadow MSJFordham MCMaxwell C. Borland HMaxwell. Of leaves can make it difficult to attain full reversion similar to the nocturnal CO2 exchange would not a! And Chinnock, 2013 ) ] and also reflects differences in the closely related species with CAM photosynthesis ( acid! ( PEP ) is not known perennial Calandrinia that dries back to the host C3-to-CAM shifts inherent constitutive., 2016. https: //biologydictionary.net/cam-plants/ across their ranges by a fully developed leaf to! Particularly promising water stress has been demonstrated in whole plants ( Winter 1974. … reversibility distinguishes facultative CAM good for then receives an electron from NADH and becomes a molecule of.! Carbon fixation evolved by some plants in the C3 mode zamiifolia ( Lodd..! Triangulare changes from C3 to CAM were performed under the conditions described in.... ( Keeley, 1983 ) state of zero nocturnal CO2 exchange for a shoot of the of. Like: oxaloacetate + NADPH + MDH → malate + NADP+ was present throughout the.. Dominated d 8, and pigment genes have been uncovered are achieved than with induction by drought will conserved! A single-cell pathway in P. oleracea ) or are believed to lack stomata ( P. grandiflora ) Calandrinia...

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