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Breeding System and Pollination Biology of Paeonia Delavayi (Peaoniaceae), an Endangered Plant in the Southwest of China

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Breeding System and Pollination Biology of Paeonia Delavayi (Peaoniaceae), an Endangered Plant in the Southwest of China Pak. J. Bot ., 46(5): 1631-1642, 2014. BREEDING SYSTEM AND POLLINATION BIOLOGY OF PAEONIA DELAVAYI (PEAONIACEAE), AN ENDANGERED PLANT IN THE SOUTHWEST OF CHINA KUI LI 1,2,3 , BAOQIANG ZHENG 1,2,3 , YAN WANG 1,2,3 * AND LIN ZHOU 1,2,3 1Research Institute of Forestry, Chinese Academy of Forestry, Beijing 100091, China 2Key Laboratory of Tree Breeding and Cultivation, State Forestry Administration, Beijing 100091, China 3State Key Laboratory of Tree Genetics and Breeding, Beijing 100091, China *Corresponding author’s e-mail: [email protected] Abstract Breeding system and pollination biology of Paeonia delavayi (Peaoniaceae) from Shangri-La, Yunnan Province, southwest of China were studied. Flowering phenologies and flower visitors were observed or collected from 2008 to 2011. The pollen viability, stigma receptivity and pollination efficiency of different visitors were detected and tested. The florescence lasted for 6- 9d in a single flower from mid-May to late June. A high percentage of flower damage promoted early anther dehiscence. Flowers started disseminating pollen at 1-2 d after flowering, and lasted for 5-6 d. Pollen viability could be preserved for more than 10 d at normal temperature. High seed rate from the stigma was observed at 1 d before flowering to 3d after flowering, and the dissemination hysteresis was defined as protogyny. The P/O ratios were 6,124 to 9,713:1, suggesting that the larger quantity of pollen to increased the seed setting rate. Three species of bees, eight species of beetles, seven species of syrphid flies, four species of ants, and three species of butterflies were observed on the flowers. P. delavayi rewarded to the visitors by releasing fragrance, providing pollen and nectar. On the bodies of the visitors under stereomicroscope and scanning electron microscope (SEM), much pollen from the plants of similar flowering period inner community were found which indicated that these incompatible visitors were not species-specific pollinators. The bagging experiments showed that P. delavayi was self- incompatible and no apomixes. Anemophily only played a minor role in the fertilization. A few seeds with poor plumpness can be produced by geitonogamy. Seed setting rate of artificial xenogamy was higher than natural pollination. Artificial control of the visitors’ species showed bees being the most important pollinators. Beetles and ants participated in pollination to some extent and were unstable. Syrphid flies and butterflies were very unreliable with low pollination efficiency. Reproductive success depended largely on cross-pollination assisted by pollinator activities, especially the bees. Key words: Ants; Beetles; Bees; Endangered plant; Flowering time; Breeding system; Paeonia delavayi ; Pollination Biology; P/O ratio; Stigma receptivity. Introduction distributed in the United States, had self-pollination characteristics. Through radioiodine labeling, Schlising Paeonia delavayi , as a plant of Paeoniaceae, Paeonia (1971) indicated that the beetles carried and disseminate and Sect. Moutan (Hong & Pan, 1999), is mainly distributed pollens among flowers of different plants. Turpin & from central to northwest Yunnan, and some in southwest Schlising (1976) conducted in-depth observational studies Sichuan and southeast Tibet. As an endemic species in and found that a total of four families, 17 species of bees, southwest of China (Wang, 1997), P. delavayi is distributed and one species of beetles participated in pollination. Based in the southernmost part under the wild tree peony group. on the research of the visiting insects on P. lactiflora in the The China Plant Red Data Book listed P. delavayi as an Gaogesitaihanwula Natural Preserve in Inner Mongolia, endangered species (Feng, 1992), and as a Grade II state Hong & Liu (2006) demonstrated that with the absence of protected plant of the Regulations of the People’s Republic apomixes and cleistogamy and with weak selfing, P. of China on Wild Plants Protection in 1996. Since 2008, we lactiflora insects played a major role in cross pollination. have conducted a large number of investigations in With regard to woody plants of Paeonia , the pollination distribution areas of P. delavayi . A total of 12 populations experiment were conducted by Luo et al ., (1998) in were found in Kunming, Dali, Lijiang, Shangri-La, Deqin, connection with P. suffruticosa subsp . and Weixi, Yunnan, as well as in Nyingchi, Tibet (Li et al ., spontanea , distributed in southern Shanxi Province, China, 2009). Based on the investigations, P. delavayi grows in was show that the species had neither apomixes nor individually or in clusters on forest edges, under secondary automatic self-pollination. In addition, the species with forests, or among bushes of mountains at elevations between unisexual flowers producing few seeds, has weak selfing 2400 and 3600 m. Human intervention has led to frequent and low seeding rate. Moreover, a total of only five species changes in the population structure of P. delavayi , and its of bees and four species of scarabs were involved in its effective seeding productivity decreased, with asexual pollination. The studies on Paeonia pollination were reproduction characteristics (Li et al ., 2012). In recent years, extremely limited. Focusing on P. delavayi , this study was the morphology, palynology, molecular biology, and seed performed as a preliminary research on the breeding and germination characteristics of P. delavayi have been biological characteristics of its pollination, including its investigated in a number of studies (Wang, 2007; Li et al ., flowering time, floral characters, pollen viability, stigma 2011). However, no detailed report on its breeding system in receptivity period and pollen-ovule ratios (P/O). This study the wild, including flowering habits, pollination ecology and aims to investigate the pollination mechanism of P. biological characteristics, etc. delavayi , its breeding system and its relationship with the At present, researches on Paeonia pollinations are reproductive biological characteristics, to provide a limited extremely. Grant (1964) reported in his theoretical basis for the effective protection and rational observational studies that the herb P. californica , mainly utilization of germplasm resources of wild P. delavayi . 1632 KUI LI ET AL ., Materials and Methods germinate after storage at 4°C for 1yr and after pollination seeded successfully, maintaining a high germination rate Study site: The study was carried out near the ski resort, at room temperature for 10 d (Lv et al., 2010). For the approximately 25 km west of Shangri-La County (27° 57' stigma receptivity, fresh pollens were obtained from the N, 99° 35' E) in Northwest of Yunnan Province, China. same population, and the pollen experiment was The area is the most large, concentrated and abundant conducted with the flower at 2 d before and 5d after the distribution region of P. delavayi , and has a temperate opening of petals. For flowers before opening of petals, highland semi-humid monsoon climate, with an altitude isolation of after emasculation was performed directly, about 3,318 m, annual average temperature of 5.9°C, had the flowers for the pollination experiment after monthly coldest average temperature of -0.4°C, monthly flowering to experience the isolation after emasculation hottest average temperature of 13.3°C, frost-free period of when the stigma was not bare and had no pollen 123.8 d, annual precipitation of 648.6 mm, annual dissemination, and with the surrounding flowers of the evaporation of 1,616.8 mm, annual sunshine time of same flowering degree as reference (Khan & Perveen, 2,155.9 h, annual average relative humidity of 69%, and 2011). The pollination time was recorded at -2, -1, 1, 2, 3, accumulated temperature above 10°C of 1,539.2°C (the 4, and 5 (from 2 d before to 5 d after flowering) in the above meteorological data was obtained from Diqing morning and afternoon, and the seeding rate was recorded Meteorological Bureau); the soil type is composed mainly when harvesting the seeds. of brown soil and limestone. Pollinators: During the flowering of P. delavayi , we Vegetation composition: The forest zone in the region is a observed various types of insect activities regularly from conifer-dominated community, and P. delavayi is 2009 to 2011 in Shangri-La of Yunnan Province. The distributed on forest edges and canyon shrub zones, with insects were captured for identification and detection of tree-layer canopy density of 0.1 to 0.2, and occasionally the pollen load by scanning electron microscopy (SEM). scattered in the understory with higher canopy density. The activity insects were observed from 8:00 am to 8:00 Moreover, the area has a shrub layer coverage of 35% to pm. A cluster about 30 plants was selected to record the 55% and herb layer coverage of 10% to 25%. The arbor, an flower-visiting frequency, movement mode, and stay time accompanying plant, composed mainly of Abies fabri , of different insects in the flowers. Pinus densata and Populus yunnanensis ; shrubs composed mainly of Salix rehderiana , Rhododendron lapponicum , R. Breeding system experiments: The following bagging pulchrum , Quercus senescens , Rosa forrestiana and experiments were performed to determine the Berberis alpine ; herbs and lianas composed mainly of characteristics of the breeding system of P. delavayi . The Clematis florida , Pteridium revolutum , Plantago asatica , flowers used were selected from regions with similar Sinopodophullum hexandrum , Iris
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