Floral and Fruit Morphology of Some Species in Garcinia Spp

ORIGINAL ARTICLE

Floral and fruit morphology of some species in Garcinia spp.

Sompong Te-chato1

Abstract Te-chato, S. Floral and fruit morphology of some species in Garcinia spp. Songklanakarin J. Sci. Technol., 2007, 29(2) : 245-252

Morphological studies of reproductive parts of some species in Garcinia revealed some differences and similarities. Flowers of these species are composed of both male and female flowers. Female flowers are solitary and occur in single or occasionally in clusters (2-10 flowers) developed at the terminal buds of young branches in case of mangosteen, but both terminal buds and axillary buds were found in the other species. Primary parts of the flowers in this genus comprised four sepals and four petals which were different in size and color. Many stamens, both filamentous and sessile anthers, are observed in both male and female flowers in different location. The number of anthers varied among the species ranging from 20 to 40 sets in phawa and approximately 135 sets in somkhag. The colors of fruits are species specific ranging from yellow (somkhag and ma-phut) to red (phawa) or deep purple (mangosteen) and different in size, shape, rind characters and flavor. Three of six species produce no seed to few seeds (1-2). Viability of pollens ranges from 100% (cha-muang, phawa and ma-dun) to 0% (mapood).

Key words : Garcinia, floral morphology, mangosteen, somkhag, phawa

1Ph.D. (Plant Cell Technology), Assoc. Prof., Department of Plant Science, Faculty of Natural Resources, Prince of Songkla University, Hat Yai, Songkhla, 90112 Thailand. Corresponding e-mail: [email protected] Received, 14 March 2006 Accepted, 25 August 2006 Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 246 Te-chato, S. ∫∑§—¥¬àÕ ¡ªÕß ‡µ™–‚µ1 ≈—°…≥–∑“ß —≥∞“π¢Õß¥Õ°·≈–º≈„πæ◊™ °ÿ≈ Garcinia ∫“ß™π‘¥ «. ß¢≈“π§√‘π∑√å «∑∑. 2550 29(2) : 245-252

°“√»÷°…“≈—°…≥–∑“ß —≥∞“π¢Õß¥Õ°·≈–º≈„π °ÿ≈ Garcinia ∫“ß™π‘¥ æ∫∑—Èß∑’Ë¡’§«“¡§≈â“¬§≈÷ß°—π·≈– ·µ°µà“ß°—π ¥Õ°¢Õßæ◊™™π‘¥¥—ß°≈à“«Õ“®‡ªìπ‡æ»ºŸâÀ√◊Õ‡æ»‡¡’¬ ¥Õ°µ—«‡¡’¬¡’≈—°…≥–‡ªìπ¥Õ°‡¥’Ë¬«Ê À√◊Õ‡ªìπ™àÕ (2-10 ¥Õ°) ‡°‘¥µ√ßª≈“¬¬Õ¥ ÿ¥ („π¡—ß§ÿ¥) À√◊Õ‡°‘¥∑—Èßª≈“¬¬Õ¥ ÿ¥ ·≈–µ“¢â“ß¢Õß°‘Ëß („π™π‘¥Õ◊Ëπ∑’Ë‡À≈◊Õ) ¥Õ° ª√–°Õ∫¥â«¬°≈’∫‡≈’È¬ß 4 °≈’∫ ·≈–°≈’∫¥Õ° 4 °≈’∫ ¡’ ’ √√·≈–¢π“¥·µ°µà“ß°—π ‡° √„π¥Õ°∑—Èß Õßª√–‡¿∑¡’ ∑—Èß∑’Ë¡’·≈–‰¡à¡’°â“π™Ÿ ¡’µ”·Àπàß∑’Ëµ—Èß·µ°µà“ß°—π ¡’‡° √µ—«ºŸâ®”π«π¡“°∑—Èß∑’Ë¡’·≈–‰¡à¡’°â“π™Ÿ ´÷Ëß®”π«ππ—Èπº—π·ª√ µ“¡ ªï™’ å µ—Èß·µà 20-40 ™ÿ¥ „πæ–«“∂÷ß 135 ™ÿ¥ „π â¡·¢° ’¢Õßº≈·µ°µà“ß°—πµ—Èß·µà ’‡À≈◊Õß („π â¡·¢° ¡–æŸ¥) ∂÷ß·¥ß (æ–«“) À√◊Õ¡à«ß‡¢â¡ (¡—ß§ÿ¥) ¢÷Èπ°—∫™π‘¥ ¢π“¥ √Ÿª√à“ß≈—°…≥–º‘«º≈ ·≈–√ ™“µ‘¢Õßº≈·µ°µà“ß°—π „π ®”π«π 6 ™π‘¥∑’Ë»÷°…“ æ∫«à“¡’ 3 ™π‘¥∑’Ë‰¡à √â“ß‡¡≈Á¥À√◊Õ √â“ß·µàπâÕ¬¡“° (1-2 ‡¡≈Á¥) §«“¡¡’™’«‘µ¢Õß‡√≥Ÿµ—Èß·µà 0% (¡–æŸ¥) ∂÷ß 100% (™–¡«ß æ–«“·≈–¡–¥—π)

1¿“§«‘™“æ◊™»“ µ√å §≥–∑√—æ¬“°√∏√√¡™“µ‘ ¡À“«‘∑¬“≈—¬ ß¢≈“π§√‘π∑√å Õ”‡¿ÕÀ“¥„À≠à ®—ßÀ«—¥ ß¢≈“ 90112

Garcinia belongs to the family Guttiferae or species of Garcinia, G. mangostana, G. malacensis Clusiaceae. The genus arose in southeast Asia, and G. hombroniana. He showed that four char- ranging from the southern part of Thailand and acters; latex and petal color, sessile stigma and peninsular Malaysia to Indonesia and to some parts fruit color of G. mangostana resembled those of of the Philippines (Richards, 1990c). However, the G. malacensis. Another three characters of G. exact origin of the genus is still debated. Accord- mangostana, stigma, stamen/staminode lobe, and ing to Whitmore (1973), the genus is listed as fruit resembled those of G. hombroniana. Based having 39 species with another ten unidentified or on these similarities he stated that G. mangostana unnamed in this region. Among these species, only is a hybrid between G. hombroniana and G. a few are known and used in daily life in Thailand, malaccensis. However, there have been no research namely G. mangostana L. (now being grown as an reports which confirm this phenomenon. orchard fruit tree) G. atroviridis Griff, G. speciosa The purpose of this study is to reveal floral Wall., G. cowa Roxb. and G. dulcis Kurz (Lim and fruit morphology and pollen viability in some et al., 1983). The latter four species possess both important species of Garcinia which have been medicinal and horticultural properties. G. found in southern Thailand for further improve- atroviridis Griff, G. dulcis Kurz and G. cowa Roxb ment of those species by conventional and bio- contain a high concentration of hydroxy citric acid technological methods. (HCA), especially in the fruit and leaf. This chemical regulates some enzymes in the hydro- Materials and methods lysis pathway in order to efficiently convert fat or fatty acid into energy (Te-chato, 1997). G. speciosa Plant materials Wall. can grow well under drought conditions. So In this investigation, six species of Garcinia it is used as a rootstock for mass propagation of namely G. mangostana L. (mangosteen), G. mangosteen. speciosa Wall. (phawa), G. atroviridis Griff. Richards (1990b) reported using morpholo- (somkhag), G. dulcis Kurz. (ma-phut), G. cowa gical characters for identification among the three Roxb. (cha-muang) and G. schomburgkiana Pierre Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 247 Te-chato, S.

(ma-dun) were collected from the Plant Science number of stained pollens divided by total number Research Station, Prince of Songkla University, of pollen in the same field multiplied by 100. Hat Yai campus and private orchards in the Narathiwat province. Reproductive parts (flowers Results and fruits) were collected for investigation. Floral and fruit morphology of Garcinia spp. Comparative study on floral and fruit mor- Morphological studies of reproductive parts phology of Garcinia spp. of various species in Garcinia revealed some Reproductive parts of each species were differences and similarities (Table 1, Figure 1). morphologically compared. Within this group, the Most wild species and male plants or male flowers six species are differentiated by 13 characters as have a long period of flowering and flower more shown in Table 1. than one time a year, whereas cultivated species, (e.g. mangosteen) bear fruit only once a year. In Pollen viability analysis the case of cultural management, off-season For testing viability of pollen, anthers from flowering can be induced. Among these species, each species of Garcinia were excised and placed mangosteen contains a concentrate yellow latex in a drop of 0.5% acetocarmine on a glass slide. affecting the quality of its fruits. The anthers were gently squeezed to release the inner pollen. The slide was covered with a cover Floral parts glass and excess staining solution blotted. The Flower slide loaded with a mixture of pollen and staining Flowers of these species are either male or solution was heat by alocohol flame. Morphology female or both which occur separately on different and viabilty of pollen from each species were plants or appear together on the same plant. Female observed and counted under inverted microscopy. flowers are solitary and occur singly or occasion- Viable pollens were scored and interpreted as the ally in clusters (2-3) as found in mangosteen,

Table 1. Floral and fruit morphological characters of some species in Garcinia.

Characters G. cowa G. speciosa G. atroviridis G. mangostana G. dulcis G. schomburgkiana Cha-muang Phawa Somkhag Mangosteen Ma-phut Ma-dun Flowering July-Sept. June-Sept., July-Sept. Mar-April, July-Sept. Jan-Mar(-June), Jan-Mar July-Sept. July-Sept. Latex Yellow White Yellow Yellow White Yellow Petal color Light yellow Cream- Purple Red Light green Pink Light yellow Stigma - Stipitate Sessile Sessile Stipitate - Stigma Surface - Smooth Corrugated Corrugated Corrugated - Stigma lobe - 20% diameter 80% diameter 20% diameter 20% diameter Stamen mass Clustered Lobed Round Isolated Lobed Clustered Female Flower - No staminodes Staminodes Staminodes Staminodes - Pollen Viability 96-100% 93-100% 3-5% 0.1-1% 0% 95-100% Fruit shape - Oval Globose Globose Globose - Fruit color - Red Green Purple Yellow - Fruit surface - Smooth Wrinkle Smooth Lobed - Fruit flavor - Astringent Sour Sweet/sour Sour -

- No character found in these species Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 248 Te-chato, S.

Figure 1. Floral morphology of Garcinia in some species (bar = 1 cm) A: G. mangostana B: G. atroviridis C: G. schomburgkiana D: G. cowa E: G. dulcis E: G. speciosa [Color figure can be viewed in the electronic version] phawa, somkhag, ma-dun and cha-muang but somkhag and ma-phut. Ma-dun and cha-muang clusters of 6-10 flowers are usually found in ma- produce flowers of nearly the same size. In case of phut. In mangosteen, the flowers develop at the color, both somkhag and ma-dun have red to dark terminal buds of young branches in case of red color while mangosteen has a pinkish red one. mangosteen; whereas in the other species, the The other species have petal color ranging from flowers develop at both terminal and axillary buds. light green to yellow (Table 1, Figure 1).

Petals and sepals Stamens and pistil Primary parts or construction of the flowers Many stamens, both filamentous and sessile in this genus are very similar. They are composed anthers, are observed in the flower. Generally, the of four sepals and four petals. Sepals of some stamens develop in both male and female flowers. species are separated into two whorles; outer and In female flowers of mangosteen, ma-phut and inner ones. Petal sizes and colors of these species somkhag, they are located on the petal around the differ. Subjective evaluation indicated that ovary at the lower position than stigma. The first mangosteen has the biggest size, followed by two species produce filamentous anthers while the Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 249 Te-chato, S.

A B

C Figure 2. Characteristics of stamen (arrows) containing in female flowers of Garcinia spp. A: G. mangostana B: G. dulcis C: G.atroviriddis

Figure 3. Male flowers with (A,B arrows) and without (C,D,E) rudimentary ovary (bar = 1 cm) A: G. speciosa B: G. atroviridis C: G. cowa D: G. schomburgkiana E: G. dulcis [Color figure can be viewed in the electronic version] latter species produces sessile ones (Figure 2a,b,c). extend over the entire ovary (Figure 3a,b). Male In the case of male flowers in phawa and somkhag, flowers of the two species also produce a small stamens develop as a tube or fused together and non-functional or rudimentary ovary. For cha- Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 250 Te-chato, S.

Figure 4. Viable pollen contained in anther of male flower staining with 0.5% carmine. A: G. speciosa B: G. cowa

Figure 5. Fruit morphology of some Garcinia species (bar = 5 cm) A: Garcinnia mangostana B: G. speciosa C: G. dulcis D: G. atroviridis [Color figure can be viewed in the electronic version] muang, ma-dun and ma-phut, the male flowers minutes. The unfertilized ovary develop to mature have only a cluster of sessile anther without a fruit within 4 to 6 months depending upon the rudimentary ovary but with viable pollens (Figure species. The colors of fruits are specific to each 3c,d,e). The number of anthers varied among the species, ranging from green in somkhag, yellow species ranging from 20 to 40 sets in phawa and in ma-phut, to pinkish red in phawa and purple in about 135 sets in somkhag. mangosteen (Figure 5). Moreover, size, shape, rind characters and flavor differ as shown in Fruit and seed Table 1 and Figure 5. Nucellar tissue in ovary also Fruits of these species were reported to develops to be apomictic seed through agamo- develop without pollination and fertilization spermy (Richards, 1990a). Number, shape, and size (Richards, 1990a). After blooming for at least 24- of the seeds varied. Among the four species which hour period, petals wither and drop within a few set fruit, phawa has the highest number of seeds Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 251 Te-chato, S.

(6-8 per fruit), while the other three species produce supporting technique promotes early growth of few seeds (1-2) to no seed. However, all of the mangosteen (Sdoodee et al., 1992). The species seeds are recalcitrant which leads to a limited that is suitable for root supporting is phawa. This distribution of the species. species is best in combating problems compared with the other species. DNA works also revealed Pollen viability analysis that phawa is very close to mangosteen (Te-chato Inside the anther, a high number of 93-100% et al., 2000). Accordingly, pollination of the two viable pollens were found in male flowers of cha- species is promising. muang, phawa and ma-dun, whereas there was no viable pollen in ma-phut (Figure 4). Very low Conclusion viable pollens were observed in mangosteen (0.1- 1%) and somkhag (3-5%). For female or pistillated Only few species of Garcinia have been flowers of all species, only 0-1% viable pollen listed in daily life of southern Thailand. At least of was found. According to this phenomenon these four species are used as fresh fruit and medicinal stamens are so called staminode. purpose. Each species has different floral morphology and pollen viability. Some have both male Discussion and female flowers while other have only male or female flowers. Data from this morphological The nature of plants in Garcinia differ from observation and pollen viability investigation species to species. Some can grow under shade might be of great importance in crossing some while the others survive under arid conditions or species which are very close e.g. mangosteen and full sunlight. Among these species, mangosteen is phawa. In future, new characters of mangosteen, very sensitive to unsuitable environments, espec- especially drought tolerance, obtained from these ially under drought, as mangosteen is a very slow crossings is possible. growing tree and seed germination is very poor. Studies on the morphology of these species are References very rare. It is reported that most or all Garcinia Lim, M., Hemapat, T., Wanachit, W. and Suwankiri, P. spp. are apomictic and the fruit develops without 1983. Survey and collection of Lansium and fertilization (Richards, 1990a). Moreover, pollin- Garcinia in Southern Thailand. IBPGR News- ation has not been reported by any researchers letter 10: 10-12. except in G. hombroniana (Richards, 1990b). From Richards, A.J. 1990a. Studies in Garcinia, dioecious this investigation, morphological characteristics tropical forest trees: agamospermy. Botanical J. are of great importance for helping us to know the of the Linnean Society 103: 233-250. way for pollination. The data of pollen viability Richards, A.J. 1990b. Studies in Garcinia, dioecious reveal the possibility of crossing among the tropical forest trees: the phenology, pollination species, especially between phawa and mangosteen, biology and fertilization of G hombroniana in order to transfer drought resistance. In the Pierre. Botanical Journal of the Linnean Society future, commercial planting of mangosteen will 103: 251-256. reduce the cost of agricultural practices. In the past, Richards, A.J. 1990c. Studies in Garcinia, dioecious grafting mangosteen on the rootstock of related tropical forest trees: the origin of the mangosteen species was applied for that purpose; however, (G. mangostana L.). Botanical Journal of the grafting success both in vitro and ex vitro was Linnean Society 103: 301-308. quite low (Te-chato et al.,1992; 1995). Recent work Sadooee, S., Lim, M. and Yong, S. 1992. Effect of root indicates that self-grafting (grafting of mangosteen supporting on growth of mangosteen (Garcinia on self-rootstock) also provides a good result mangostana). Songklanakarin J. Sci. Technol. (Sdoodee et al., 1992). On the other hand, root 14: 327-335. Songklanakarin J. Sci. Technol. Floral and fruit morphology of some species in Garcinia spp. Vol.29 No.2 Mar. - Apr. 2007 252 Te-chato, S.

Te-chato, S. 1997. Isolation and culture of protoplast of and Genomic Technology for Crop Improvement somkhag (Garcinia atroviridis Griff.) to micro- (Eds. K. Oono, T. Komatsuta, K. Kadowaki and colony. Songklanakarin J. Sci. Technol. 19: 255- D. Vaughan), pp. 63-68, Tsukuba: Sato Printing 262. Co., Ltd. Te-chato, S., Lim, M. and Chumroenruk, M. 1995. Te-chato, S., Lim, S. and Kulsiroratana, J. 1992. In vitro Grafting vitro-shoot of mangosteen on rootstock micrografting of mangosteen. Songklanakarin of pawa ex vitro. Songklanakarin J. Sci. Technol. J. Sci. Technol. 14: 353-359. 17: 245-250. Whitmore, T.C. 1973. Garcinia L. In Tree flora of Te-chato, S., Lim, M. and Masahiro, M. 2000. Diversity Malaya (ed. T.C. Whitmore) vol. 2, pp 196-225. of Garcinia spp. and interspecies relationships Kuala Lumpur: Forest Department, Ministry of by DNA analyses. In Integretion of Biodiversity Primary Industries, Malaysia.