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Intraspecific Variation in Floral Organs and Structure in Brassica Rapa L. Analyzed by Principal Component Analysis

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Intraspecific Variation in Floral Organs and Structure in Brassica Rapa L. Analyzed by Principal Component Analysis Breeding Science 56 : 189–194 (2006) Note Intraspecific Variation in Floral Organs and Structure in Brassica rapa L. analyzed by Principal Component Analysis Syafaruddin1), Yosuke Yoshioka1), Atsushi Horisaki2), Satoshi Niikura2) and Ryo Ohsawa*1) 1) Graduate School of Life and Environmental Sciences, University of Tsukuba, 1-1-1 Tennodai, Tsukuba, Ibaraki 305-8572, Japan 2) Tohoku Seed Company, 1625 Nishihara, Himuro, Utsunomiya, Tochigi 321-3232, Japan Key Words: anther-stigma separation, floral characteristics, F1 seed production. Many biologists have been interested in the evolution incompatible parental lines are planted and pollination is of floral organs as adaptations for effective animal pollina- performed by wild insects, such as honeybees, bumblebees tion. Much of their interest has focused on the effect of floral and hoverflies. The seed yields of F1 hybrids depend on the morphology on the efficiency of pollen removal and deposi- pollinators’ flights among individuals of different parental tion during pollinator visits (Campbell 1989, Murcia 1990, lines, and on the pollination efficiency of individual flowers Young and Stanton 1990, Nishihiro et al. 2000, Yang et al. during pollinator visits. Therefore, to improve F1 seed pro- 2002, Kudo 2003), and the attractiveness to a pollinator for duction, a detailed investigation of floral organs and struc- which there is a high probability that pollen will be placed ture is required. The objective of the present study was to where fertilizing a conspecific ovule is likely to occur investigate the variability of floral characteristics in a wide (Schemske and Bradshaw 1999). Many investigators have range of B. rapa cultivars. indicated significant effects of the characteristics of floral We used four inbred parental lines of some F1 hybrids, organs and structures, especially the spatial relationship be- which become genetically homogeneous through five to tween stigma and anthers, on the flowers’ pollination effi- seven generations of self-crossing, and 30 cultivars that rep- ciencies, in both self-fertilizing and outcrossing species resented high genetic variability across eight varietal groups (Holtsford 1992, Conner et al. 1995, Karron et al. 1997, of B. rapa, in order to investigate intraspecific variation in Motten and Stone 2000, Elle and Hare 2002). In other stud- floral characteristics (Table 1). The plants were grown in a ies, it was reported that the floral size and number play an glasshouse at the Agriculture and Forestry Research Center, important role in attracting pollinating insects (Andersson University of Tsukuba (Tsukuba, Japan), from November 1996, Conner and Rush 1996), such as bees that will use the 2002 until May 2003. We chose three plants per line or cul- flower shape as a visual cue in foraging (Gegear and Laverty tivar, and sampled three flowers from the main stem per 2001, Giurfa and Lehrer 2001). plant (306 flowers in total). The flowers that had newly From a plant breeder’s point of view, floral morpholo- opened on each sampling day were used. gy should be considered to have the potential to increase or After removal of the four petals, each flower was decrease seed production in both self-fertilizing and out- placed sideways on a stage, and its image was projected onto crossing crops. In fact, in many investigations of floral or- a computer display via a Keyence digital microscope (VH- gans, it was showed that the floral morphology influenced 5000, Keyence Co., Osaka, Japan). We measured eight char- the pollination efficiency (Levin et al. 1994, Uga et al. acteristics of the floral organs — long stamen length (LSL), 2003a, 2003b, Yoshioka et al. 2005), and that a lower degree short stamen length (SSL), long anther length (LAL), short of stigma exsertion increased the rate of self-fertilization anther length (SAL), ovary length (OL), stigma height (SH), (Namai et al. 1992, Yashiro et al. 2001, Syafaruddin et al. stigma width (SW) and style length (SL) (Fig. 1) — using 2002, Kobayashi et al. 2004). In crops that outcross by in- image analysis software (VH-Analyzer, Keyence Co.). sect pollination, it remains to be determined what kind of To summarize the information about the eight charac- floral characteristics attract pollinating insects and, in partic- teristics of the floral organs, we performed a principal com- ular, what kind of floral morphology is advantageous to pol- ponent analysis (PCA) based on a variance–covariance ma- lination efficiency. trix. The scores of the principal components (PC) were used In seed-production fields of Brassica rapa L., self- in the analysis to identify the characteristics responsible for the majority of the variation in the floral structure. To deter- Communicated by Y. Takahata mine the effect of each PC, we reconstructed the floral struc- Received June 17, 2005. Accepted January 11, 2006. ture from the eight variables, reverse-calculated by letting *Corresponding author (e-mail: [email protected]) the score for a certain PC be equal to the mean ± 2 standard 190 Syafaruddin, Yoshioka, Horisaki, Niikura and Ohsawa Table 1. List of lines and cultivars across eight varietal groups used in the present study and mean values of eight floral characteristics. SE and CV denote the standard error of the mean and coefficient of variation, respectively Line no. Cultivar name Varietal group LSL SSL LAL SAL OL SH SW SL LVC-01 Chirimen-hakusai pekinensis 0.72 0.51 0.20 0.21 0.46 0.06 0.11 0.16 LVC-02 Nagasaki-hakusai pekinensis 0.78 0.58 0.20 0.20 0.58 0.07 0.10 0.15 LVC-03 Hankekyuu-santosai pekinensis 0.89 0.66 0.21 0.20 0.44 0.06 0.10 0.15 LVC-04 Maruba-santosai pekinensis 0.78 0.60 0.21 0.19 0.53 0.08 0.11 0.26 LVC-05 Bansei-osakashirona pekinensis 0.79 0.58 0.20 0.18 0.44 0.08 0.11 0.19 LVC-06 Banseimana pekinensis 0.78 0.59 0.19 0.17 0.46 0.08 0.11 0.26 LVC-07 Hiroshimana pekinensis 0.82 0.61 0.21 0.20 0.47 0.06 0.11 0.20 LVC-08 Nabana campestris 0.85 0.61 0.21 0.19 0.53 0.08 0.11 0.21 LVC-09 Seppakutaina chinensis 0.82 0.62 0.19 0.18 0.51 0.07 0.12 0.21 LVC-10 Nagaokana chinensis 0.77 0.57 0.21 0.24 0.47 0.08 0.12 0.25 LVC-11 Shigatsu-shirona chinensis 0.75 0.55 0.21 0.18 0.42 0.07 0.09 0.16 LVC-12 Shinobu-fuyuna rapifera 0.82 0.61 0.21 0.19 0.57 0.08 0.09 0.26 LVC-13 Niigata-tona campestris 0.90 0.61 0.22 0.22 0.59 0.09 0.13 0.27 LVC-14 Sendai-yukina chinensis 0.79 0.60 0.21 0.20 0.51 0.07 0.10 0.21 LVC-15 Kukitachina campestris 0.99 0.76 0.28 0.26 0.60 0.07 0.10 0.26 LVC-16 Katsuyama-mizuna campestris 1.25 0.96 0.32 0.32 0.78 0.07 0.11 0.26 LVC-17 Kumamoto-kyona rapifera 0.90 0.65 0.23 0.22 0.52 0.08 0.11 0.27 LVC-18 Osakina rapifera 0.80 0.61 0.25 0.24 0.43 0.07 0.12 0.30 LVC-19 Wase-aburana campestris 0.88 0.66 0.24 0.23 0.53 0.09 0.13 0.29 LVC-20 Shinshu-yukina campestris 0.93 0.67 0.28 0.27 0.58 0.07 0.11 0.29 LVC-21 Shizuoka-kyona campestris 0.84 0.63 0.25 0.22 0.52 0.07 0.09 0.18 LVC-22 Tatsai narinosa 0.79 0.56 0.17 0.17 0.42 0.08 0.11 0.12 LVC-23 Chugoku-saishin parachinensis 0.90 0.66 0.19 0.19 0.62 0.08 0.13 0.19 LVC-24 Kotsaitai chinensis 0.90 0.68 0.23 0.22 0.58 0.08 0.12 0.21 LVC-25 Pakchoi chinensis 0.90 0.76 0.23 0.23 0.58 0.09 0.13 0.22 LVC-26 Sensuji-kyomizuna japonica 0.89 0.61 0.23 0.22 0.67 0.07 0.09 0.22 LVC-27 Mibuna japonica 0.84 0.63 0.28 0.29 0.57 0.07 0.10 0.23 LVC-28 Nozawana rapa 0.87 0.63 0.27 0.27 0.53 0.07 0.09 0.21 LVC-29 Gensuke-kabuna rapa 0.85 0.62 0.26 0.25 0.45 0.06 0.10 0.23 LVC-35 Kamokabu rapa 0.78 0.57 0.22 0.21 0.46 0.06 0.11 0.17 HaO-017 pekinensis 0.67 0.41 0.20 0.20 0.45 0.07 0.12 0.18 HaO-251 pekinensis 0.72 0.48 0.21 0.21 0.46 0.08 0.13 0.21 HaO-263 pekinensis 0.84 0.60 0.24 0.23 0.42 0.07 0.11 0.21 HaO-314 pekinensis 0.85 0.59 0.23 0.22 0.44 0.08 0.10 0.18 Total Mean 0.84 0.62 0.23 0.22 0.52 0.07 0.11 0.22 SE (10−1) 0.17 0.15 0.06 0.06 0.14 0.01 0.02 0.08 CV 0.12 0.14 0.14 0.16 0.16 0.12 0.11 0.21 LSL: long stamen length (cm), SSL: short stamen length (cm), LAL: long anther length (cm), SAL: short anther length (cm), OL: ovary length (cm), SH: stigma height (cm), SW: stigma width (cm), SL: style length (cm).
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