Autogamy and Allogamy in Genus Lathyrus
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Lathyrus Lathyrism Newsletter 2 (2001) Autogamy and allogamy in genus pots, 20 cm in diameter, the plants were grown under homogenous conditions (25°C and 12h /day length) in Lathyrus an experimental garden, at the National Agronomic Research Institute of Tunisia (Upper Semi-arid). Plant flowering occurred from April (L. cicera, L. sativus and L. aphaca) to June (L. latifolius and L. sylvestris). N. Ben Brahim1, D. Combes2 and M. 3 Three types of treatments were performed: selfing by Marrakchi bagging flowers in order to prevent insect visitation of the flower (Treatment A), natural pollination without 1. Laboratoire de botanique, Institut National de la castration of anthers (Treatment B) and natural Recherche Agronomique de Tunisie, pollination following castration of anthers (Treatment Rue Hédi Karray, 2049, Ariana, Tunisie. C). Each type of operation was performed on 15 2. Laboratoire d’Ecologie Moléculaire, IBEAS, flowers per plant, 5 plants were studied per species. Université de Pau et des Pays de Ladours, France. Two fertility parameters were measured for each plant 3. Laboratoire de Génétique et de Biologie on 15 flowers per plant: the frequency of flowers Moléculaire, Faculté des Sciences de Tunis. giving pods (p/f) and the mean number of seeds per pod (g/p). Email : (1) [email protected] The values of the ratio p/f ranged from 0 to 1, the transformation Introduction f(x) = arc sin√x, where x = p/f, was used. In Tunisia the genus Lathyrus is represented by 15 (11) native species , distributed essentially in marginal Statistical analysis. The data of the fertility parameters environments in the north of the country. Many of were analysed by analysis of variance to test them are of considerable agronomic interest as forage differences between species and between species and (L. ochrus and L. articulatus) and as human food (L. breeding system effects. The Duncan test (7) was used cicera and L. sativus). These species are under utilised to compare means. The SAS procedure ANOVA was and in the past little effort has been made to expand the used. selection and breeding of Lathyrus as a food crop. However, many Lathyrus species have potential as an alternative pulse in many cropping systems as they are Results tolerant to low rainfall conditions. Research is now being conducted to address aspects of improvement of Selfing by bagging flowers. The three perennial some species such as L. sativus, including reducing species L. latifolius, L. sylvestris and L. tuberosus did concentrations of the neurotoxin β-N-oxalyl-L-α, β- not develop pods by selfing, the ratios p/f and g/p were diaminopropanoic acid (ODAP) in the seed, both zero (Table 2) and indicate that these species are introducing insect and disease resistance, and strictly outcrossing. For that reason they were excluded increasing yield (4). from statistical analysis. The variance analysis showed highly significant differences between annual species In order to identify useful material for agriculture and for the two studied fertility parameters: number of for breeding programs, we studied breeding systems flowers giving pods p/f (F ratio 6.04) and number of and analysed some aspects of floral biology of fourteen seeds per pod g/p (F ratio 37.41). annual and perennial Lathyrus species. The comparison of means using Duncan test at 5 % Materials and methods (Table 3) reveals a regrouping of most species for high Fourteen annual and perennial Lathyrus species means for the number of flowers giving pods (p/f). belonging to four sections according to Kupicha’s However, L. odoratus is distinguished from other classification (10) were analysed, each species was species by a low mean for p/f (0.25) (Table 2). represented by one population. All these species are diploid (2n=2x=14). Studied species, their biological The comparison of means at 5% showed that L. type and their sites of origin are listed in Table 1, only nissolia had the highest number of seeds per pod (9.30) L. sativus, L. cicera and L. odoratus (the ornamental (Tables 2 and 4). This can be explained partly by the sweetpea) are cultivated. high number of ovules per ovary in this species (Table 5). L. aphaca, L. cicera, L. sativus and L. setifolius had The seeds used in this study were germinated in petri the lowest means for g/p (range 2.44 to 4.12). dishes on wet filter paper. They were manually scarified in order to increase their germination percentage. Seedlings were transplanted in October to 21 Lathyrus Lathyrism Newsletter 2 (2001) Table 1. Lathyrus species studied, nomenclature of Kupicha(10). Species Abbreviation Section Habit Origin L. annuus AN Lathyrus Annual Tunisia L. aphaca A Aphaca Annual Italy L. articulatus AR Clymenum Annual Tunisia L. cicera C Lathyrus Annual Tunisia L. hirsutus H Lathyrus Annual Italy L. latifolius L Lathyrus Perennial Tunisia L. nissolia N Nissolia Annual France L. odoratus O Lathyrus Annual Tunisia L. ochrus OC Clymenum Annual Tunisia L. sativus SA Lathyrus Annual Tunisia L. setifolius SE Lathyrus Annual Italy L. sylvestris S Lathyrus Perennial France L. tingitanus TI Lathyrus Annual Portugal L. tuberosus T Lathyrus Perennial Hungary Table 2. Frequency of flowers giving pods and mean number of seeds per pods in selfing and in natural pollination. Species Frequency of Mean number of flowers giving seeds per pod pods (p/f) (g/p) ABCABC L. annuus AN 0.82 0.84 0 5.33 5.43 0 L. aphaca A 0.72 0.79 0 4.12 4.32 0 L. articulatus AR 0.64 0.88 0 5.31 5.38 0 L. cicera C 0.91 0.86 0.13 3.58 3.44 3 L. hirsutus H 0.88 0.90 0 5.80 6.25 0 L. latifolius L 0 0.85 0.11 0 6.34 6.37 L. nissolia N 0.75 0.82 0 9.30 9.75 0 L. ochrus OC 0.81 0.80 0 5.11 4.63 0 L. odoratus O 0.25 0.81 0.20 5.84 6.67 5.46 L. sativus SA 0.53 0.80 0.09 2.82 2.50 2.57 L. setifolius SE 0.66 0.72 0 2.44 2.70 0 L. sylvestris S 0 0.73 0.13 0.20 5.07 6.40 L. tingitanus TI 0.68 0.89 0 6.33 6.70 0 L. tuberosus T 00.880040 A selfing by bagging B natural pollination without castration C natural pollination following castration However, L. nissolia is characterized by high values of Natural pollination without castration of anthers. g/p. L. setifolius and L. sativus have the lowest ratio of Variance analysis indicated no significant differences g/p and the lowest number of ovules per ovary (Table for the ratio p/f (F ratio1.56) and highly significant 5). differences for fertility parameter g/p (F ratio 48.35). In natural pollination without castration of anthers, the The comparison of means by Duncan test at 5 % (Table values of the two fertility parameters are higher than 4) demonstrated a regrouping between all species for those observed in selfing with bagging anthers (Table the number of pods per flower (p/f) (Table 3). 2). This is particularly prominent for the three 22 Lathyrus Lathyrism Newsletter 2 (2001) perennial species L. latifolius, L. sylvestris and L. with castration than in natural pollination without tuberosus , and also L. odoratus, all of which did not castration. Some of this reduction could be attributed to develop pods in selfing. The values of p/f increase lesion caused by castration. respectively from 0 to 0.88 for L. tuberosus, from 0 to Comparison of species by breeding system effects. 0.85 for L. latifolius, from 0 to 0.73 for L. sylvestris Variance analysis with two classification criteria and from 0.25 to 0.81 for L. odoratus. indicated that there is a species by breeding system effect, which is highly significant for the two studied Natural pollination following castration of anthers. parameters (F ratio 9.75, for p/f and F ratio 47.23 for The results of natural pollination following castration g/p). revealed that the castrated flowers from annual species (L. aphaca, L. nissolia, L. articulatus, L. ochrus, L. The comparison of averages reflects a regrouping for tingitanus, L. annuus, L. setifolius and L. hirsutus) did species for the parameter p/f. However, a heterogeneity not develop pods and seeds (Table 2). However, for is observed for the ratio g/p: L. nissolia is characterised five species (L. odoratus, L. sativus, L. latifolius, L. by the highest means, while L. setifolius and L. sativus sylvestris and L. cicera) the ablation of anthers caused have the lowest means (Table 4). a clear reduction in the number, but did not completely inhibit the development of pods and seeds. This Comparison between selfing with bagging, natural indicates clearly that the three cultivated annual species pollination without castration of anthers and (L. odoratus, L. sativus and L. cicera) can be pollinated natural pollination following castration of anthers. either by their own pollen or by foreign pollen. The comparison of the three breeding systems using Duncan test at 5 % reveals a separation of breeding Variance analysis revealed no significant differences systems for the two parameters p/f and g/p. For the between the five species for the parameter p/f (F ratio ratio g/p, the two breeding systems: natural pollination 2.58) (Table 3) and highly significant differences for and selfing by bagging are grouped with the highest g/p (F ratio 12.83) (Table 4). The comparison of the means.