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(Passiflora Edulis Sims) and Fruits and Cladodes of the Yellow Pitaya [Hylocereus Megalanthus (K

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(Passiflora Edulis Sims) and Fruits and Cladodes of the Yellow Pitaya [Hylocereus Megalanthus (K CROP PHYSIOLOGY Stomatal behavior in fruits and leaves of the purple passion fruit (Passiflora edulis Sims) and fruits and cladodes of the yellow pitaya [Hylocereus megalanthus (K. Schum. ex Vaupel) Ralf Bauer] Comportamiento estomático en frutos y hojas de gulupa (Passiflora edulis Sims) y en frutos y cladodios de pitahaya amarilla [Hylocereus megalanthus (K. Schum. ex Vaupel) Ralf Bauer] Camilo Sánchez1, Gerhard Fischer1, and Danny Wilson Sanjuanelo2 ABSTRACT RESUMEN Plants as C3 and CAM react photosynthetically different but Las plantas C3 y CAM reaccionan fotosintéticamente dife- both can grow in the same agroecological zone in the tropics. rentes, sin embargo, ambas pueden crecer en la misma zona Therefore we studied the behavior of stomatal opening in fruits agroecológica en los trópicos. Por esta razón se evaluó el com- and leaves of the purple passion fruit and fruits and cladodes portamiento de estomas en frutos y hojas de gulupa y en frutos of the yellow pitaya was studied under natural growing condi- y cladodios de pitahaya amarilla bajo condiciones naturales tions in Granada and Fusagasuga, Cundinamarca (Colombia). en los municipios de Granada y Fusagasugá, departamento Imprints were made on the surface of leaves, fruits and cladodes de Cundinamarca (Colombia). Se hicieron improntas sobre using cosmetic enamel impressions. Three cycles were carried la superficie de las hojas, frutos y cladodios. Se realizaron tres out, each cycle took 72 hours, obtaining three different samples lecturas, cada una duró 72 horas, haciendo impresiones cada every 3 hours; then the impressions were observed by micro- 3 horas sobre tres muestras diferentes para luego observarlas scope and the opened and closed stomata were counted in each en microscopio, haciendo un conteo de estomas abiertos y cer- species. In each sampling, data of solar radiation, temperature rados en cada especie. En cada muestreo se obtuvieron datos and relative humidity (RH) were measured. The purple passion de radiación solar, temperatura y humedad relativa (HR). En fruit had the typical behavior of a C3 plant in the leaves as well gulupa se encontró un comportamiento típico de una planta as the fruits, and a positive correlation between the stomatal C3 tanto para las hojas como para los frutos, además de encon- aperture and radiation and temperature was found, along with trar una correlación positiva entre la apertura estomática y la a negative correlation between stomatal aperture and RH. The radiación y temperatura, así como una correlación negativa pitaya showed the typical behavior of a CAM plant with a nega- entre la apertura estomática y la HR. En pitahaya se encontró tive correlation between the stomatal opening and radiation and un comportamiento típico de una planta CAM, observando temperature, as well as a positive correlation between stomatal una correlación negativa entre la apertura estomática y la opening and RH. Radiation, temperature and RH affected the radiación y temperatura, así como una correlación positiva stomatal opening in the fruits and cladodes. Stomatal densities entre apertura estomática y HR. La radiación, la temperatura differed greatly between the species and plant organs. In the y la HR afectaron la apertura estomático en frutos y cladodios. purple passion fruit, 106.53 stomata per mm2 leaf surface were Las densidades estomáticas contrastaron altamente entre las found, but only 12.64 stomata per mm2 fruit surface; whereas especies y los órganos de la planta. En gulupa fueron encontra- in the pitaya, 11.28 and 1.43 stomata per mm2 were found on dos 106,53 estomas/mm2 en la superficie de las hojas, pero solo the cladodes and fruits, respectively. 12,64 estomas/mm2 en la del fruto; mientras que en pitahaya fueron encontrados 11,28 y 1,43 estomas/mm2 en cladodios y frutos, respectivamente. Key words: tropical fruits, C3 plants, CAM plants, stomata, Palabras clave: frutas tropicales, plantas C3, plantas CAM, stomatal movement. estomas, movimiento estomático. Introduction by more flowering species than probably any other plant The purple passion fruit Passiflora( edulis Sims) is a vigor- family (Menzel and Simpson, 1994). It is an exotic fruit ous woody perennial vine which belongs to the Passiflora- from the region between the Brazilian Amazon, the north ceae family, which includes 550 species and is represented of Argentina and Paraguay (Morton, 1987). In Colombia, Received for publication: 3 December, 2012. Accepted for publication: 29 March, 2013. 1 Department of Agronomy, Faculty of Agronomy, Universidad Nacional de Colombia. Bogota (Colombia). [email protected] 2 Faculty of Science and Technology, Universidad de Ciencias Aplicadas y Ambientales (UDCA). Bogota (Colombia). Agronomía Colombiana 31(1), 38-47, 2013 the production area of the purple passion fruit was esti- key role in the economy of water in plants, since more than mated by Agronet (2012) for 2011 at 520 ha, principally in 90% of plant water is lost through leaf stomata (Hess, 2004). the departments of Antioquia, Boyacá, Cundinamarca, When stomata are closed, the main pathway for water loss Huila, Quindío and Santander. These areas are located is through cuticles (Fageria et al., 2006). at altitudes between 1,800 and 2,400 m, with a medium temperature range of 15-20°C (Jiménez et al., 2012) and Wong et al. (1978) described three mechanisms which 1,500-2,200 mm of rainfall per year (Fischer et al., 2009). cause stomatal response to light. First, the guard cells seem to possess receptive pigments of light that directly detect The pitaya or dragonfruit Hylocereus[ megalanthus (K. changes in light intensity and change the stomatal conduc- Schum. ex Vaupel) Ralf Bauer], from the tropical Americas, tance accordingly. Second, the guard cells can indirectly was introduced to Colombia in the mid 1980s due to the feel a change in the intensity of light through a change in interest of Japanese experts and the Federación Nacional the partial pressure of intercellular CO2 (Ci) that is caused de Cafeteros de Colombia. Production in Colombia is con- by a change in the photosynthetic rate of the mesophyll centrated in the departments of Boyacá, Huila, Santander, tissue. Third, the guard cells can indirectly respond to Valle del Cauca, Cundinamarca, Tolima, Caldas, Bolivar, changes in the rate of photosynthesis by a chemical trans- and Quindío, reaching an area in 2011 of 691 ha (Agronet, missible signal that is transported from the mesophyll cells 2012). The best cultivating areas are located at altitudes to the guard cells. In many species, stomata show a diurnal between 1,500 and 1,850 m, with a medium temperature rhythm, opening by day and closing by night, with light range of 18-21°C and 1,500-2,000 mm of rainfall per year stimulating stomatal opening, with blue wavelengths being (Corredor, 2012). most effective (Öpik and Rolfe, 2005). Stomata are formed in the epidermis of all plant organs, Within normal ranges (10-25°C), changes in temperature except the roots (Kramer, 1989), preferentially on the leaves. have little effect on stomatal behavior (Evert and Eichhorn, Stomata are small pores or valves that control the entry of 2012), but temperatures higher than 30 to 35°C can lead to carbon dioxide assimilated in photosynthesis and optimize stomatal closure which can be due to an indirect response water use efficiency, essential for plant survive and perfor- from the water stress or from an increase in the respira- mance (Bergmann and Sack, 2007). Adjusting the number tory rate that induces an increase in the CO2 inside the leaf or movements (opening and closing) of stomata is essential (Salisbury and Ross, 1992). for regulating the water use of crop plants (Pritchard and Amthor, 2005). As specialized structures in the epidermis, Stomata regulate water loss (Pospílová and Dodd, 2005), they are formed by two elongated guard cells enclosing a and stomatal closure responds to the decrease of the mois- pore between them, across which plants absorb CO2 and ture in the environment (Kearns and Assmann, 1993). Mott liberate O2 to the environment (Hetherington and Wood- and Parkhurst (1991) suggested that a stomatal closing in ward, 2003). Each stoma is surrounded by two guard (oc- response to increasing water stress involves a diminished clusive) cells, which change their shape to open and close availability of water for the guard cells and thus the turgor the pores (Evert and Eichhorn, 2012). The apertures are so decreases. The apoplastic abscisic acid (ABA) concentration small that at most 3% of the total leaf surface is occupied around the guard cells is crucial in determining stomatal by the pores (Öpik and Rolfe, 2005). responses; when roots are stressed through water deficit, ABA travels by the xylem to the leaves, signaling stomata The stoma opens due to water absorption by the guard closure (Pospílová and Dodd, 2005). cells (Assmann, 1993; Wang et al., 1998), which in turn is caused by an increase in the solute; potassium ions moving In almost all plants, low concentrations of CO2 cause the towards the occlusive cells as the stoma opens (Salisbury opening of stomata, even in darkness (Kramer, 1989); if and Ross, 1992). This behavior is the result of interac- CO2 circulates freely through the leaves, they will open tions between physiological factors and environmental more. On the contrary, a high concentration of CO2 in conditions (Assmann and Wang, 2001; Hetherington and the leaves can provoke stomatal closing, in the presence of Woodward, 2003). light or darkness (Salisbury and Ross, 1992; Kearns and Assmann, 1993). Environmental factors that affect stomatal movement include light, temperature and CO2 concentration (Evert The number of stomata varies greatly (Kramer, 1989). and Eichhorn, 2012). Also, the stomatal response plays a Lake et al. (2001) determined that stomatal density (or Sánchez, Fischer, and Sanjuanelo: Stomatal behavior in fruits and leaves of the purple passion fruit (Passiflora edulis Sims) and fruits and cladodes..
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