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Crassulacean Acid Metabolism in the Gesneriaceae Lonnie J

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Crassulacean Acid Metabolism in the Gesneriaceae Lonnie J Roger Williams University DOCS@RWU Feinstein College of Arts & Sciences Faculty Feinstein College of Arts and Sciences Publications 1986 Crassulacean acid metabolism in the Gesneriaceae Lonnie J. Guralnick Roger Williams University, [email protected] Irwin P. Ting Elizabeth M. Lord Follow this and additional works at: http://docs.rwu.edu/fcas_fp Part of the Arts and Humanities Commons Recommended Citation Guralnick LJ, Irwin. P Ting, Elizabeth M. Lord. 1986. "Crassulacean acid metabolism in the Gesneriaceae." American Journal of Botany 73 (3):336-345 This Article is brought to you for free and open access by the Feinstein College of Arts and Sciences at DOCS@RWU. It has been accepted for inclusion in Feinstein College of Arts & Sciences Faculty Publications by an authorized administrator of DOCS@RWU. For more information, please contact [email protected]. Crassulacean Acid Metabolism in the Gesneriaceae Author(s): Lonnie J. Guralnick, Irwin P. Ting and Elizabeth M. Lord Source: American Journal of Botany, Vol. 73, No. 3 (Mar., 1986), pp. 336-345 Published by: Botanical Society of America, Inc. Stable URL: http://www.jstor.org/stable/2444076 Accessed: 18-10-2017 14:40 UTC REFERENCES Linked references are available on JSTOR for this article: http://www.jstor.org/stable/2444076?seq=1&cid=pdf-reference#references_tab_contents You may need to log in to JSTOR to access the linked references. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at http://about.jstor.org/terms Botanical Society of America, Inc. is collaborating with JSTOR to digitize, preserve and extend access to American Journal of Botany This content downloaded from 198.7.243.146 on Wed, 18 Oct 2017 14:40:38 UTC All use subject to http://about.jstor.org/terms Amer. J. Bot. 73(3): 336-345. 1986. CRASSULACEAN ACID METABOLISM IN THE GESNERIACEAE' LoNNIE J. GURALNICK, IRWIN P. TING, AND ELIZABETH M. LORD Department of Botany and Plant Sciences, University of California, Riverside, California 92521 ABSTRACT The occurrence of the Crassulacean acid metabolism (CAM) was studied in four epiphytic species of the Gesneriaceae: two neotropical species, Codonanthe crassifolia and Columnea linearis, and two paleotropical species, Aoschynanthus pulcher and Saintpaulia ionantha. Gas exchange parameters, enzymology, and leaf anatomy, including mesophyll succulence and rel- ative percent of the mesophyll volume occupied by airspace, were studied for each species. Codonanthe crassifolia was the only species to show nocturnal CO2 uptake and a diurnal organic acid fluctuation. According to these results, Codonanthe crassifolia shows CAM-cycling under well-watered conditions and when subjected to drought, it switches to CAM-idling. Other characteristics, such as leaf anatomy, mesophyll succulence, and PEP carboxylase and NADP malic enzyme activity, indicate attributes of the CAM pathway. All other species tested showed C3 photosynthesis. The most C3-like species is Columnea linearis, according to the criteria tested in this investigation. The other two species show mesophyll succulence and relative percent of the leaf volume occupied by airspace within the CAM range, but no other characters of the CAM pathway. The leaf structure of certain genera of the Gesneriaceae and of the genus Peperomia in the Piperaceae are similar, both having an upper succulent, multiple epidermis, a medium palisade of one or a few cell layers, and a lower, succulent spongy parenchyma not too unlike CAM photosynthetic tissue. We report ecophysiological similarities between these two distantly related families. Thus, the occurrence of CAM-cycling may be more common among epiphytic species than is currently known. THE CRASSULACEAN ACID METABOLISM (CAM) - Both are similar to typical CAM metabolism, pathway of photosynthesis is regarded as an but show slight modifications in their gas ex- adaptation to arid and semiarid environments change parameters. CAM-cycling is character- (Kluge and Ting, 1978). CAM is characterized ized by C3-type gas exchange accompanied by by nocturnal CO2 uptake and malic acid ac- a diurnal acid fluctuation. Stomatal conduc- cumulation. Subsequent daytime stomatal clo- tance is high during the day and low at night, sure and decarboxylation of malic acid pro- but there is still a nocturnal acidification pri- duces CO2 as the substrate for photosynthesis, marily due to recycling of respiratory CO2. The resulting in less water loss under periods of second modification, called CAM-idling, usu- high evaporative demand. The leaves of CAM ally occurs when CAM or CAM-cycling species plants are also characterized by tightly packed are water stressed. Exogenous gas exchange is chlorenchyma with large vacuoles where the halted due to stomatal closure, but a slight malic acid accumulates (Gibson, 1982). In ad- diurnal acid fluctuation is still observed. dition, the mesophyll is generally not differ- Not all CAM species are found in arid en- entiated into palisade and spongy parenchyma vironments; in fact, many occur in xeric mi- layers (Kluge and Ting, 1978). These charac- croenvironments of mesic regions, such as rock teristics of CAM are important under water- outcrops (Martin and Zee, 1983), and others limiting conditions. are tropical epiphytes that periodically become Two new modifications of CAM have re- drought stressed (Winter et al., 1983; Sinclair, cently been reported (Kluge and Ting, 1978). 1984; Ting et al., 1985). The CAM pathway of photosynthesis occurs in tropical epiphytes ' Received for publication 9 May 1985; revision ac- of the Bromeliaceae (McWilliams, 1970; Me- cepted 25 September 1985. dina and Troughton, 1974), Orchidaceae We thank Kathleen Eckard for her help with the leaf (Knauft and Arditti, 1969; McWilliams, 1970; sections and Bill Crone for his assistance in the stereology. We thank Janet Hann and Alfredo Huerta for their help Neales and Hew, 1975; Sinclair, 1984), Cac- with the HPLC. Special thanks to Jan Lippert for doing taceae (Wiehler, 1982), Rubiaceae (Winter et many acid titrations and also the graphics. We thank Patti al., 1983), and the Piperaceae (Sternberg, De- Garcia for typing the many revisions of the manuscript. niro and Ting 1984; Sipes and Ting, 1985). Supported in part by grants from the National Science Foundation, PCM 82-00366, DMB-8416981, and PCM CAM is also found in some epiphytic ferns of 8303-6 19. the Polypodiaceae (Wong and Hew, 1976). 336 This content downloaded from 198.7.243.146 on Wed, 18 Oct 2017 14:40:38 UTC All use subject to http://about.jstor.org/terms March, 1986] GURALNICK ET AL.-CRASSULACEAN ACID METABOLISM 337 The largely tropical Gesneriaceae has about ative percent of the leaf volume occupied by 120 genera and 3,000 species, of which 20% airspace ? SE. are epiphytic in their habitat. A number of the Mesophyll succulence (Sm) was calculated epiphytic gesneriads overlap in range with Pep- from the ratio of total tissue water to total eromia spp. of the Piperaceae and, in addition, chlorophyll (Kluge and Ting, 1978) in fresh they both share a similar upper succulent mul- leaf samples. Tissue water content was deter- tiple epidermis (Ting et al., 1985). Since it was mined by drying samples in a microwave oven recently reported that the genus Peperomia until dry and subtracting the dry weight from shows attributes of CAM (Ting et al., 1985), the fresh weight. Total chlorophyll was mea- we investigated some epiphytic Gesneriaceae sured by grinding fresh leaf tissue in 80% ace- for the occurrence of CAM. In this paper, we tone and then centrifuging at 1,500 g. A 1 ml report that the Gesneriaceae has epiphytic aliquot was added to 4 ml 80% acetone and species showing attributes of CAM. The in- then measured spectrophotometrically (Ar- vestigation included a comparison of physio- non, 1949). Sm was determined for the whole logical and anatomical characters of CAM and leaf. Results are expressed as g H20 mg- Chl ? C3-related gesneriad species. We wanted to as- SE. certain which anatomical characters might cor- relate to the CAM pathway. We report here Titratable acidity-Three leaves per time our investigations of the gas exchange param- point were collected, weighed, and frozen until eters, leaf anatomy, and enzymology of four assayed. Samples were ground in glass distilled gesneriads: two neotropical and two paleo- water with a tissue grinder and titrated with tropical species. 0.01 N KOH to a pH 7.0 endpoint. Data are expressed as Aeq g-I FW ? SE. MATERIALS AND METHODS-Plants-Saint- paulia ionantha (H. Wendl.) plants were a gift Enzyme assays-Leaf samples were collect- from Craig Yoshida of Sunnyside Nursery of ed in triplicate, weighed, and 1 g FW was ground Hayward, Calif. Aeschynanthus pulcher (G. in a tissue grinder in 10 ml of extraction me- Don) plants were purchased from Tiegs Nurs- dium at 4 C, containing 100 mm Hepes-NaOH, ery of Riverside, California. Codonanthe cras- 10 mM MgCl2, 10 mm DTT, and 1% PVP 360 sifolia (Focke) and Columnea linearis (Oerst.) adjusted to pH 7.8 for extraction of PEP car- specimens were collected at La Selva, Costa boxylase and to pH 7.2 for NADP malic en- Rica, and propagated in glasshouses at the Univ. zyme. The sample was centrifuged at 4,000 of Calif., Riverside. Plants were propagated rpm at 4 C for 10 min, and a small aliquot was from cuttings and grown in a glasshouse under passed through a Sephadex G-25 column, which natural photoperiods during the course of the had been pre-equilibrated with the adjusted year. The glasshouse had a mean annual high Hepes-NaOH buffer.
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