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Durio Zibethinus Murray TROPICS Vol. 14 (3) Issued March 15, 2005 Phenological characteristics of reproduction and seed formation in Durio zibethinus Murray 1) 2) 2) 3) Kazuharu OGAWA *, Ahmad Makmom ABDULLAH , Muhamad AWANG and Akio FURUKAWA 1) Laboratory of Forest Ecology and Physiology, Graduate School of Bioagricultural Sciences, Nagoya University, Nagoya 464-8601, Japan 2) Department of Environmental Sciences, Universiti Putra Malaysia, 43400 UPM Serdang, Malaysia 3) Department of Biological Sciences, Faculty of Science, Nara Women’s University, Nara 630-8506, Japan *) Author to whom correspondence should be addressed. Tel: +81-52-789 4071, Fax: +81-52-789-5014, E-mail: [email protected] ABSTRACT Phenological characteristics during the reproductive processes of flower budding, flowering, fruiting and seed formation were investigated in a Durio zibethinus Murray tree growing in an experimental field of Universiti Putra Malaysia (UPM) in Selangor, Peninsular Malaysia. The survival curve showed that the relative fall rate of reproductive organs was lower at the mixed stages of flower budding, flowering and fruiting than at the early fruiting stage. The fall of flower buds commenced after the flower budding was completed. However, the falls of flowers and fruits occurred within a few days after the formation of flowering and fruiting. The periods of flower budding, flowering and fruiting were 34, 19 and 28 days, respectively. The fruit weight growth was approximated by a logistic equation, where the intrinsic growth rate varied from 0.419 to 0.794 week-1. The forming seeds germinated or possessed germination ability at the fruit growth stage where the fruit weight reached the asymptote of the logistic growth curve. Total numbers of emerged flower buds, flowers and fruits over the whole reproductive period were 1629, 1467 and 1412 per tree, respectively. The percentage of fruits with germination ability was estimated to be 1.2% of the number of fruiting ones. Key words: Durio zibethinus Murray, fruit growth, germination ability, reproductive processes, survival curve. INTRODUCTION During the life cycle of woody species, phenological behaviors are represented by sequential processes of reproduction, such as flower budding, flowering and fruiting (Harper 1977). Among the reproductive processes, the fruiting process is directly related to the processes of seed formation and germination. The seeds develop so as to possess the germination ability during the fruit growth. Clarifying whether or not forming seeds possess germination ability should provide important information on the dynamics of forest ecosystems, including natural regeneration. Phenological information on reproductive processes in tropical trees is limited, and only qualitative information, such as seasonality, is available (Longman and Jenik 1987; Whitmore 1984). Moreover, there is little information on ripening during fruit growth. To minimize measurement difficulties, a tropical tree, Durio zibethinus Murray (clone D99B) grown in the orchard was chosen. D. zibethinus is valued for its fruits, as well as its timber (Idris 1990; Subhadrabandhu et al. 1991; Smith et al. 1992; Yaacob and Subhadrabandhu 1995). Since wild fruit trees are known to be as productive as their descendants developed in orchards (Whitmore 1984), the present results on D. zibethinus grown in an orchard could be applied to the phenology of forest trees of this species. The measurement methods were designed to determine the survival curves of each reproductive process: flower budding, flowering and fruiting. After fruiting, the fruit growth stage (where the forming seeds possess the germination ability) was determined by fitting the logistic function to the fruit weight growth curve. MATERIALS AND METHODS Sample tree This study was conducted on a 13-year-old (as of 1993) Durio zibethinus Murray tree (clone D99B) at an experimental field station of Universiti Putra Malaysia (UPM) in Selangor, Peninsular Malaysia. One tree was sampled. The tree height and stem diameter at breast height of the sample tree were 8 m and 34.1 cm, respectively. Survival curve The reproductive processes were divided into flower budding, flowering and fruiting (Fig. 1). To determine the survival 222 Kazuharu OGAWA, Ahmad Makmom ABDULLAH, Muhamad AWANG and Akio FURUKAWA curves during the reproductive processes, it was necessary to identify whether the attached and fallen Flower budding organs were flower buds, flowers or fruits. Then, all flower buds of the sample tree were marked and Flower buds n information on whether or not the petals fell, styles n 1 turned brown or ovaries became large., was recorded. Fall Un-flowering Flowering Furthermore, fruits were classified as mature or Flowers immature. n-n 1 The attached and fallen numbers of flower buds, n 2 Fall flowers and fruits were recorded 27 times at 1 to 13 day Un-fruiting Fruiting intervals from September 19, 1993 to February 12, 1994. Fruits n-n 1-n 2 The girths of branch bases were measured in October n 3 1993. There was one replication. Fall Un-maturing Maturing Fruit growth Mature fruits n-n 1-n 2 -n 3 The study tree commenced fruiting in early October n-n 1-n 2 -n 3 Fall 1993. On November 23, the lateral diameter (DS, cm) and longitudinal diameter (DL, cm) of 35 attached fruits were Fig. 1. Schematic representation of the reproductive 15 processes of flower budding, flowering and un- labeled and measured by Vernier calipers (Type CD- flowering, and fruiting and un-fruiting, together and 30, Mitsutoyo, Japan). The measurement was with maturing and immaturing. nn and with repeated for 26, 20 and 19 attached fruits on December 4, subscript stand for the flower budding number 16 and 27, 1993, respectively and 18 attached fruits on and the fallen number of reproductive organs, respectively. January 7, 1994. The number of sample fruits decreased, because this species drops fruits continuously after fruiting. The fallen fruits were collected to measure the fruit size and check the germination ability of seeds. The dry mass (w, g -1 1.45 2 fruit ) of attached and fallen fruits was estimated from the allometric equation, w=0.105 (DS DL) (R =0.98). This equation was formulated from the observed data on 192 fruits (Ogawa et al. 1996). Germination test We collected seeds from 18 fallen fruits of different fruit ripening stages from November 23, 1993 to February 12, 1994 and made the germination test (Table 1). After measuring the fresh weight of seeds, the seeds were sowed in fiberglass pots containing forest topsoil and stored on the rooftop of a building of UPM. RESULTS Relationship between the number of flower buds and branch diameter The number of branches that produced flower buds was 13; the other 13 branches did not produce flower buds. The number of emerged flower buds, ranging from 0 to 371 per branch, positively correlated with branch diameter (P<0.05) (Fig. 2). One can infer from this positive correlation that flower budding depends on materials stored in branches and current photosynthates by leaves. Survival curve Mixed process Figure 3 is a time line showing the number of attached reproductive organs (N) per tree in the processes of flower budding, flowering and fruiting on the semi-log coordinates. The absolute value of gradient of the straight line after the complement ∇ ∇ ∇ 1 N∇ of flower budding stands for the relative fall rate RFR＝－― ― during a given time interval (t ) (Ogawa et al. （ Nt） 1995). While RFR was lower at the mixed stage of flower buds, flowers and fruits than at the early fruit stage, RFR became Phenology and seed formation in Durio zibethinus 223 Table 1. Fruit dry weight at the fallen time, the number and fresh weight of sample seeds used for germination test, and germination ratio during the period of November 23, 1993 to February 12, 1994. Fruit No. Fallen time Fruit weight Sample seed Germination ratio Number Fresh weight per seed [g d. wt] [g f. wt] [%] 1 Nov. 23-Dec. 2 62.5 - - 0 2 -"- 19.6 - - 0 3 -"- 31.0 - - 0 4 -"- 20.2 - - 0 5 Dec. 2-Dec. 4 10.4 - - 0 6 -"- 11.0 - - 0 7 -"- 21.8 - - 0 8 Dec. 4-Dec. 13 127.6 - - 0 9 -"- 19.5 - - 0 10 -"- 127.6 - - 0 11 Dec. 16-Dec. 27 43.8 - - 0 12 -"- 45.5 - - 0 13 -"- 74.6 - - 0 14 Jan. 19 594.6 6 27.7 100 15 Feb. 1 386.1 7 20.1 100 16 Feb. 2 571.4 6 26.6 100 17 Feb. 12 303.3 4 21.2 100 18 Feb. 12 439.6 3 28.6 100 400 350 300 250 200 150 100 Number of flower buds per branch 50 0 0246810121416 Branch diameter [cm] Fig. 2. Relationship between the emerged number of flower buds per branch and the diameter of branch base. The correlation coefficient is 0.467 (P <0.05). 224 Kazuharu OGAWA, Ahmad Makmom ABDULLAH, Muhamad AWANG and Akio FURUKAWA 10000 Fig. 3. Survival curves in the mixed process of flower budding, flowering and fruiting on N the semi-log coordinates. ○, flower bud; □, flower bud, flower and fruit; ◇, flower and fruit; △, fruit. 1000 100 Number of reproductive organs per tree 10 Sept. Oct. Nov. Dec. Jan. Feb. Month low at the later fruit stage. Separated process Figure 4 shows a time line depicting the number of reproductive organs per tree in the separate processes of flower budding, flowering and fruiting on the semi-log coordinates. While the survival curves in the flower budding and fruiting processes showed a peak, the survival curve in the flowering process fluctuated. From the emergence curves, the periods of flower budding, flowering and fruiting were determined to be 34, 19 and 28 days, respectively. The fall and emergence curves indicate that the fall of flower buds commenced after flower budding was complete. On the other hand, the fall of flowers and fruits occurred within a few days after flowering and fruiting.
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