Water Relations of Tropical Epiphytes: I. RELATIONSHIPS BETWEEN STOMATAL RESISTANCE, RELATIVE WATER CONTENT AND THE COMPONENTS OF WATER POTENTIAL

The water relations of five species of tropical vascular epiphytesnative to Malaysia were studied. The species were ferns: Pyrrosiaadnascens (Forst.) Ching. and Pyrrosia angustata (Sw.) Ching.;orchids: Eria velutina Lindl., Dendrobium tortile Lindl. andDendrobium crumenatum Sw. Leaf resistance as a function of leafwater potential was measured for the two ferns. The criticalwater potential at which stomata closed was found to be highin each case; –0.75 MPa and –0.5 MPa respectively.The components of water potential were estimated with the pressurechamber as functions of relative water content. For each speciescell sap was found to be dilute, pressure potential low at fullturgor, and the change in relative water content between fullturgor and wilting point small. Small values of solute potentialat full turgor were also found for the ferns and E. velutinausing a vapour pressure osmometer. Values of the bulk modulusof elasticity of the leaf tissue for each species lay withinthe range of published data. The significance of these resultsfor the epiphytic way of life is discussed. Key words: Water potential, Epiphytes, Diffusive resistance, Orchid, Fern     

Water Relations of Tropical Epiphytes: II. PERFORMANCE DURING DROUGHTING

Relative water content R, water potential and leaf diffusiveconductance C₅ were monitored while five tropical epiphyteswere subjected to an extended period of drought. The speciesstudied were: ferns Pyrrosia adnascens (Forst.) Ching and Pyrrosiaangustata (Sw.) Ching, family Polypodiaceae; orchids: Eria velutinaLindl., Dendrobium tortile Lindl. and Dendrobium crumenatumSw. The ferns reached zero turgor rapidly and after only small declinesin relative water content (R) and . Beyond this point stomatalmovement seemed strongly suppressed, but the leaves continuedto lose water vapour until very low values of R were reached.Nevertheless, on re-watering water potential (), R and diffusiveconductance (C₅) returned to pre-stress levels within 3 d. The orchids showed a more gradual decline in R and , stomatalactivity was not so strongly suppressed, and night opening ofstomata was observed under stress. The relationship between and R was found for each species,curves being fitted to the data points by non-linear regression.From these analyses it was concluded that these species hadvery dilute cell sap, and consequently the change in for agiven decline in R was smaller than for most other species recordedin the literature. Key words: Orchid, Fern, Drought, Relative water content, Diffusive conductance, Water potential     

广西蕨类植物分布新记录

报道广西蕨类植物分布新记录8种,即细毛碗蕨、黑柄铁角蕨、湖南黔蕨、双胞耳蕨、台湾耳蕨、柔软耳蕨、尾叶石韦和龙骨星蕨。列出这些种的标本引证、地理分布和分类特征等。     

Structure and Histochemistry of Stomata and Epidermal Cells in Five Species of Polypodiaceae

The epidermal structure of the five species of ferns, Arthromeriswallichiana (Spr.) Ching., Drymoglossum piloselloides (Prest.),Drynaria quercifolia (L.) J. Smith, Lepisorus nudus (Hook.)Ching. and Pyrrosia nuda (Gies.) Ching., has been investigated.Fifteen types of stomatal structures have been identified ofwhich copolo-desmocytic and coperi-desmocytic are new types.Four more possible stomatal structures: ccpolo-peri-, codesmo-polo-,codesmo-peri- and duplodesmocytic, are suggested. Localizationof starch, insoluble polysaccharides, protein and lipids hasbeen examined histochemically in the guard cells, subsidiarycells and epidermal cells. In Drynaria starch plastids and plastidscontaining both starch and protein are present in guard cells.Starch plastids are present in the subsidiary cells of all speciesexcept in Arthromeris, whereas, they are present in epidermalcells of only Drymoglossum and Lepisorus. Granular or amorphousinsoluble polysaccharides (other than starch) are present inguard cells of all the species, in the subsidiary cells of Arthromeris,Drynaria and Pyrrosia, and in the epidermal cells of Pyrrosia.Except in Pyrrosia lipids are present in the guard cells. Subsidiarycells of Drynaria and the epidermal cells of Arthromeris andDrynaria show lipid bodies. The presence of plasmodesmata andectodesmata is demonstrated in the epidermal cells of Drymoglossum.     

Crassulacean acid metabolism in the epiphytic ferns Drymoglossum piloselloides and Pyrrosia longifolia: studies on responses to environmental signals

Abstract The paper reports the results of the comprehensive study of crassulacean acid metabolism in two epiphytic tropical ferns, Drymoglossum piloselloides and Pyrrosia longifolia. The plants were investigated under different light, temperature and water status. It was found that both species are obligate CAM plants. The diurnal acidity rhythm is due to the fluctuation in malic acid concentration, which accounts for the change in titratable acidity. Besides malic acid, shikimate and oxalate are found to be present, but not contributing to the CAM acid rhythm. The diurnal rhythm of malic acid content results in a corresponding rhythm in leaf water relations. Both Φ_Φ and Φ_total, were lowest at the end of the night, i.e. when the level of malic acid was highest. The effects of temperature on CO₂ exchange were inverse to those observed in other CAM plants. In both ferns studied, dark CO₂ fixation increased when the night temperature was increased. Increase in day temperature reduced CO₂ uptake during phase IV and during the following night. The observed responses of the ferns to temperature changes suggest that the in situ environmental conditions are optimal for their CAM performance. In weak light, the plants showed net CO₂ output during the midday deacidification period. Increases in light intensity reduced such CO₂ output. Under drought conditions, the CO₂ exchange in the ferns was reduced to zero within 5–6 d, indicating that the ferns studied are more susceptible to water deficiency than other CAM plants. This could be due to a higher cuticular conductance for water. The results are discussed, in particular, in relation to CAM performance of epiphytes growing in the wet tropics.     

Relationships among rainfall,leaf hydrenchyma,and Crassulacean acid metabolism in Pyrrosia lanceolata (L.) Fraw. (Polypodiaceae) in central Taiwan

Leaf succulence is common among drought-adapted plants, including many tropical and subtropical epiphytic species. A prominent anatomical feature of many such succulent leaves is a clear, water-storing tissue often referred to as “hydrenchyma” (water-storage parenchyma). Functionally, hydrenchyma appears to store water for use by the leaf during drought. Although this has been confirmed in several laboratory studies, field studies linking the amount of hydrenchyma in plants with availability of water in their environment are lacking. In this study, the relative amount of leaf hydrenchyma in one of the most widely distributed epiphytes in Taiwan, Pyrrosia lanceolata, was measured in plants growing along a gradient of annual mean precipitation from 2048 to 3688 mm. In addition, because Pyrrosia lanceolata is a Crassulacean acid metabolism (CAM) plant, the amount of CAM activity was also examined in plants along the gradient. At each of seven sites along the precipitation gradient, leaves were collected, and, using thin mid-leaf slices, the relative areas of the leaf cross-sections occupied by hydrenchyma were determined. CAM, measured as diel changes in leaf acidity, was measured in plants from each site in the field, after 3 days of water-saturation in the greenhouse, and also after 14 days without water in the greenhouse. Regressions of relative hydrenchyma with ten environmental variables in the dry season revealed that the amount of hydrenchyma was significantly and positively correlated with monthly mean number of rainless days, monthly mean number of days with daily mean temperature over 30 °C, and monthly mean temperature. During the wet season, relative hydrenchyma area correlated only with the amount of cloud cover, and the correlation was negative. All plants at all sites exhibited CAM acid fluctuations in the field, under water-saturated conditions, and after desiccation. The largest nocturnal acid accumulations were found when plants were well-hydrated in the field and in the greenhouse, although evidence of drought-induced elevations of CAM was found at the drier sites. The results of this study indicate that the amount of leaf hydrenchyma was greatest in areas with warmer, drier environments. Also, drought-induced elevation of CAM activity occurred in plants from drier sites. This may help to explain the wide range of environments inhabited by this epiphytic fern in Taiwan.     

In situ Studies of Crassulacean Acid Metobolism in Drymoglossum piloselloides, an Epiphytic Fern of the Humid Tropics

This paper reports autecological field-studies in Singaporeon Drymoglossum piloselloides (L.) Presl., an epiphytic fernof the humid tropics which is capable of performing Crassulaceanacid metabolism (CAM). As indicated by the gas exchange patternsand by the occurrence of a diurnal malic acid rhythm, the plantalso features CAM in situ at its natural sites. Both in well-wateredand in naturally droughted plants external CO₂ was taken upsolely during the night. Water stress decreased nocturnal CO₂uptake,but left the synthesis and storage of malic acid unaffected.This indicates that CO₂ recycling of respiratory CO₂ by CAMis ecophysiologically important at the high night temperaturestypical of the tropical habitats of the fern. The plants showeda diel fluctuation of cell-sap osmotic pressure which paralleledthat of malic acid, while the fluctuation of the xylem tensionfollowed the curve of transpiration more closely than it followedthat of the malic acid content. CAM in D. piloselloides wasclearly not limited by natural access to mineral ions and nitrogen.It is concluded that the ecophysiological advantage of CAM forD. piloselloides lies in a better water use efficiency as comparedwith C₃ ferns and in the salvaging of carbon by CO₂ recycling. Key words: CAM, epiphytic ferns, gas exchange, water relations     

Crassulacean acid metabolism in the ZZ plant, Zamioculcas zamiifolia (Araceae)

Zamioculcas zamiifolia (Araceae), a terrestrial East African aroid, with two defining attributes of crassulacean acid metabolism (CAM) (net CO(2) uptake in the dark and diel fluctuations of titratable acidity) is the only CAM plant described within the Araceae, a mainly tropical taxon that contains the second largest number of epiphytes of any vascular plant family. Within the Alismatales, the order to which the Araceae belong, Z. zamiifolia is the only documented nonaquatic CAM species. Zamioculcas zamiifolia has weak CAM that is upregulated in response to water stress. In well-watered plants, day-night fluctuations in titratable acidity were 2.5 μmol H(+)·(g fresh mass)(-1), and net CO(2) uptake in the dark contributed less than 1% to daily carbon gain. Following 10 d of water stress, net CO(2) uptake in the light fell 94% and net CO(2) uptake in the dark increased 7.5-fold, such that its contribution increased to 19% of daily carbon gain. Following rewatering, dark CO(2) uptake returned to within 5% of prestressed levels. We postulate that CAM assists survival of Z. zamiifolia by reducing water loss and maintaining carbon gain during seasonal droughts characteristic of its natural habitat.     

FOLIAR TRICHOMES OF PLEUROTHALLIDINAE (ORCHIDACEAE): FUNCTIONAL SIGNIFICANCE

Foliar surfaces of nine pleurothallid orchids (Cryptophoranthus lepidotus L. O. Wms., Octomeria sp., Pleurothallis pidax Luer, P. poeppigii Lindl., P. revoluta (Ruiz & Pav.) Garay, Restrepiella ophiocephala (Lindl.) Garay & Dunsterv., Restrepia muscifera Rchb. f. ex Lindl., Scaphosepalum rapax Luer, Stelis endresii Rchb. f.), four nonpleurothallid orchids (Cyrtopodium punctatum (L.) Lindl., Encyclia cochleata (L.) Lemee, E. tampensis (Lindl.) Small, Paphiope-dilum hybrid) and five tillandsioid bromeliads (Catopsis nutans (Sw.) Griseb., Tillandsia fas-ciculata Sw., T. streptophylla Scheidw., T. stricta Soland., T. tectorum E. Morr.) were assayed for permeability to Ca, S and P ions. Compared to leaves of the atmospheric bromeliads, those of the orchids proved less permeable to Ca and S. Moisture exchange profiles were determined for two of the bromeliads and three orchids; bromeliad leaves were able to rehydrate completely, orchid leaves only partially. Absorptive capacities of trichomes borne by five taxa were tested by autoradiographic analysis. Unlike bromeliad trichomes, those of the orchids exhibited no capacity to accumulate ³H-leucine. These results and other considerations described here suggest that, unlike tillandsioid bromeliads, neither the pleurothallid nor the nonpleurothallid orchids examined in this survey rely heavily on shoots for mineral and moisture procurement.     

山西蕨类植物新资料

报道了山西省新记录的蕨类植物6种1变种,隶属6科、7属,它们是井栏边草、东方狗脊、贵阳铁角蕨、雅致针毛蕨、沼泽蕨、毛轴假蹄盖蕨和高大耳蕨.     

Relationship between fern development and CAM in Pyrrosia piloselloides (L.) Price

Pyrrosia piloselloides (L.) Price is a constitutive CAM plant in the sporophytic phase of its life-cycle. Newly developed sporophytes, still attached to the gametophytes, showed signs of CAM expression in terms of diurnal changes in titratable acidity of the tissues. The gametophytes did not exhibit CAM.     

Regeneration of drought-stressed gametophytes of the epiphytic fern, Pyrrosia pilosellodes (L.) Price

The epiphytic habitat represents a highly dynamic environment, and water deficit is one of the common factors that affects growth and development of epiphytes. Gametophytes of the epiphytic fern, Pyrrosia piloselloides (L.) Price, were able to tolerate up to 50 days of drought. Upon rehydration, cells that recovered from water stress were capable of forming new gametophytes. The ability of gametophytes to recover from desiccation plays an important role in the survival and growth of the fern species under natural conditions. Received: 20 May 1998 / Revision received: 5 August 1998 / Accepted: 21 August 1998     

中国兰科植物研究杂记(英文)

报道了中国兰科植物一新记录种互生对叶兰[Neottia alternifolia(KingPantl.)Szlach.];并对紫婉石斛(Dendrobium transparens Wall.ex Lindl.)的形态特征、生境等进行了描述。紫婉石斛与兜唇石斛[Dendrobium aphyllum(Roxb.)C.E.Fischer]相似,区别在于唇瓣中央具深紫红色大斑块,唇瓣正面具柔毛;互生对叶兰是鸟巢兰属自养类型种类中唯一具有两片互生叶的种类,极易同该属其他种类区分。     

Relationship between fern development and CAM in Pyrrosia piloselloides (L.) Price

Pyrrosia piloselloides (L.) Price is a constitutive CAM plant in the sporophytic phase of its life-cycle. Newly developed sporophytes, still attached to the gametophytes, showed signs of CAM expression in terms of diurnal changes in titratable acidity of the tissues. The gametophytes did not exhibit CAM. This revised version was published online in June 2006 with corrections to the Cover Date.     

Root and Shoot Growth of Plants Treated with Abscisic Acid

Young seedlings of Capsicum annum L., Commelina communis L.and maize (Zea mays L.) were subjected to a mild water-stressingtreatment and/or treated with abscisic acid (ABA). Plants rootedin soil received a soil-drying treatment and their leaves weresprayed with a 10–⁴ M solution of ABA. Plants grown insolution culture were stressed by the addition of polyethyleneglycol (PEG) to the rooting medium and ABA was also added tothe rooting medium, either with or without PEG. The effectsof both treatments on the growth of roots and shoots and theultimate root: shoot dry weight ratio were very similar. Shootgrowth was limited both by water stress and by ABA application;while there was some evidence that mild water stress and/orABA application may have resulted in a stimulation of root growth.More severe water stress reduced the growth of roots but theoverall effect of stress was to increase the ratio of rootsto shoots. Capsicum annum L., Commelina communis L., Zea mays L., water stress, abscisic acid     

Bacteria associated with the roots of epiphytic orchids

This work is the first to report the isolation and identification of bacteria colonizing the roots of the tropical epiphytic orchids Acampe papillosa (Lindl.) Lindl. and Dendrobium moschatum (Buch.-Ham.) Swartz. and bacteria inhabiting inner layers of the aerial and substrate roots of A. papillosa. We showed by the example of this epiphyte that associative bacteria are present in large amounts on the aerial but not the substrate roots. We isolated and identified bacteria from the substrate roots of D. Moschatum and from its growth substrate (pine bark). The structure of the intercellular matrix of the associative bacteria was studied.Translated from Mikrobiologiya, Vol. 73, No. 6, 2004, pp. 825–831.Original Russian Text Copyright © 2004 by Tsavkelova, Cherdyntseva, Netrusov.     

湖南省石松类和蕨类植物分布新记录

在2004年出版的《湖南植物志》中收录有标本考证的湖南省石松类和蕨类植物683种41变种10变型[1],截至2014年底,又陆续发表了70余个新记录[2-9]。2013年出版的《Flora of China》结合现代分子系统学和最新的研究成果,对大量植物物种进行归并和科属的重新划分,作为湖南省优势科属资源的一些种类数量也相应减少,如水龙骨科( Polypodiaceae)的鳞果星蕨属( Lepidomicrosorium Ching et K. H. Shing )和盾蕨属( Neolepisorus Ching ),鳞毛蕨科( Dryopteridaceae)的复叶耳蕨属( Arachniodes Blume)以及以武陵山区为中心分布的黔蕨属( Phanerophlebiopsis Ching)(归并到复叶耳蕨属Arachniodes Blume)等,书中仅记录湖南省分布的石松类和蕨类植物约434种(含变种、亚种和杂种)[10],数量相对较少,而湖南省分布的一些较珍稀的蕨类植物未有罗列,如川黔肠蕨也Diplaziopsis cavaleriana ( Christ) C. Chr.页、中华双扇蕨( Dipteris chinensis Christ )、全缘燕尾蕨也Cheiropleuria integrifolia ( D. C. Eaton ex Hook.) M. Kato页、睫毛蕨也Pleurosoriopsis makinoi ( Maxim. ex Makino) Fomin页[1]和心叶薄唇蕨也Leptochilus cantoniensis ( Baker) Ching页等[11]。作者根据近十年的相关文献[1-9,12],对湖南省分布的石松类和蕨类植物进行了梳理,统计出631种27变种2亚种和2杂种。自2013年开始,作者对湖南省东南部的莽山、西部的小溪以及西北部的八大公山等多个国家级自然保护区进行了野外考察,发现了湖南省石松类和蕨类植物新记录6种,分属于5科6属,它们的生境及形态特征见图1。凭证标本保存于上海辰山植物标本馆( CSH )。其中一些种类分布的海拔较低,对于湖南省蕨类植物区系特点、科属结构以及物种多样性演化等研究具有潜在价值。     

海南兰科植物新资料

报道了中国兰科Orchidaceae植物一新记录属和3个中国新记录种及5个海南新记录种。其中小囊兰属Micropera Lindl．、红花小囊兰Microperapoilanei（Guill.）Garay、疏花羊耳蒜Liparis sparsiflora Aver.和美丽云叶兰Nephelaphyllum pulchrum Bl.为中国新记录;平卧曲唇兰Panisea cavalerei Schltr．、云南曲唇兰Panisea yunnanensis S．C．Chen＆Z．H．Tsi、束花石斛Dendrobium chrysanthum Wallichex Lindl．、滇南翻唇兰Hetaeria rubens （Lindl．）Benth．ex J．D．Hook．f和毛叶芋兰Nervilia plicata（Andrews）Schltr．为海南新记录种。     

Salicylic Acid Levels in Thermogenic and Non-Thermogenic Plants

The natural trigger for heat production in the thermogenic inflorescencesof Sauromatum guttatum Schott (voodoo lily) was recently identifiedas salicylic acid (SA), which induced heat production at levelsas low as 13 ng g f. wt^–1. Since then the levels of SAwere determined in other thermogenic and non-thermogenic plantspecies. In thermogenic inflorescences of five aroid species,and in male cones of at least four thermogenic cycads SA levelsduring heat production exceeded 1 µg g f. wt^–1.SA was not detected in the thermogenic flowers of a water lily,Victoria regia Lindl. (Nymphaeaceae), and Bactris major Jacq.(Palmae). Levels of salicylic acid varied substantially in thefloral parts of seven non-thermogenic species and in the leavesof 27 non-thermogenic species. Amorphophallus campanulatus Blume ex Decne, Arum italicum Mill., Arum dioscoridis Sibth. & Son., Philodendron selloum Koch, Monstera deliciosa Liebm., Encephalartosferox Bertol. f., Encephalartos hildebrandtii A. Br. & Bouché, Encephalartos gratus Prain, Dioon edule Lindl. cv. edule, Dioon edule Lindl. cv angustifolium, Sauromatum guttatum Schott, voodoo lily, Victoria regia Lindl., Bactris major Jack, salicylic acid, thermogenicity, heat production     

Induction and Repression of CAM in Sedum telephium L. in Response to Photoperiod and Water Stress

Lee, H. S. J. and Griffiths, H. 1987. Induction and repressionof CAM in Sedurn relephluni L. in response to photopcnod andwater stress.—J. exp. Bot. 38: 834–841. The introduction and repression of CAM in Sedurn telephiunmL, a temperate succulent, was investigated in watered, progressivelydrouglited and rewatered plants in growth chambers. Measurementswere made of water vapour and CO₂ exchange, titratable acidity(TA) and xylem sap tension. Effects of photoperiod were alsostudied. CAM was induced by drought under long or short days,although when watered no CAM activity was expressed. C₃-CAM intermediate plants were used for the investigation ofwater supply. Those which had received water and those drought-stressedboth displayed a similar nocturnal increase in TA, with a day-nightmaximum (H+) of 69 µmol g^–1 fr. wt. The wateredplants took up CO₂ at a maximum rate of 2?2 µmol m^–2s^–1 only in the light period, while the droughted plantsshowed a maximum nocturnal CO₂ uptake rate of 0?69 µmolm^–2 s^–1. Subsequently, as CAM was repressed, thewatered S. telephiwn displayed little variation in TA, withconstant levels at 42 µmol g^–1 fr. wt. (day 10).After 10 d of drought stress, the CAM characteristics of S.telephiurn were aLso affected, with reduced net CO₂ uptake andH+. The transition between C₃ and CAM in S. telephium can be describedas a progression in terms of the proportion of respiratory CO₂which is recycled and refixed at night as malic acid, in comparisonwith net CO₂ uptake. Recycling increased from 20% (day 1) to44% (day 10) as a result of the drought stress and was highin both the CAM-C₃ stage (no net CO2 uptake at night) and alsoin the drought-stressed CAM stage (reduced net CO₂ uptake atnight). The complete C₃-CAM transition occurred in less than8 d, and the stages could be characterized by xylem sap tensionmeasurements: CAM = 0?50 MPa C₃-CAM = 0?36 MPa C₃ = 0?29 MPa. Key words: CAM, Sedum telephium L., recycling