Flower, seed, and fruit development in three Tunisian species of Polygonum: Implications for their taxonomy and evolution of distyly in Polygonaceae

Autoři: Maher Mahmoudi aff001;  Fayçal Boughalleb aff002;  Giuseppe Pellegrino aff003;  Raoudha Abdellaoui aff002;  Nizar Nasri aff001
Působiště autorů: Université de Tunis El-Manar, Faculté des Sciences de Tunis, Tunis, Tunisia aff001;  Laboratoire des écosystèmes pastoraux et valorisation des plantes spontanées et des microorganismes associés, Institut des Régions Arides, Médenine, Tunisia aff002;  Department of Biology, Ecology and Hearth Sciences, University of Calabria, Rende (CS), Italy aff003
Vyšlo v časopise: PLoS ONE 15(1)
Kategorie: Research Article
prolekare.web.journal.doi_sk: https://doi.org/10.1371/journal.pone.0227099


Polygonum is the largest genus of Polygonaceae and 5 species are reported in Tunisia. In order to characterized flower, seed, and fruit development in Polygonum, flower and fruit of Polygonium equisetiforme (var. graecum and peyerinhoffi), P. aviculare and P. maritimum, collected from Tunisia, were examined. Flowers are composed of five oblong tepals. P. equisetiforme and P. aviculare have whitish-pink distylous flowers with dimorphism of style, filament and anther height, pollen diameter and stigma size. In contrast, P. maritimum shows white homostylous flowers. The floral vasculature showed that the tepals are inserted in one whorl and their traces arise independently in 3+2 manner. The eight stamens are arranged in a 5+3 manner and the staminal bundles arise independently in the two whorls. The epidermis and endothecium cells width were higher in P. maritimum and the lowest endothecium width was observed in P. aviculare. Polygonum aviculare and P. equisetiforme showed circular pollen with shallow colpi and trilobite pollen shape with deep colpi, while P. maritimum rarely showed shallow colpi. The ovule is anatropous with basal placentation in P. equisetiforme and P. aviculare and apical placentation in P. maritimum. The young seed coat was formed by an endotesta with thick-walled cells, a mesotesta and exotesta with thin-walled cells and a tegmen composed of radially elongated cells. The fruits of the studied species are trigonous with ovate-lanceolate shape. In P. aviculare, the exocarp is thicker compared to the two other species, in P. equisetiforme, the mature exocarp consists of smaller rectangular cells with narrow cavities, and in P. maritimum showed a thinner exocarpIn conclusion, P. equisetiforme and P. aviculare are a typically distylous species from the morphological point of view and we discussed the significance of heterostyly in Polygonaceae. From this first morpho-anatomical study of Polygonum species in North Africa, we can conclude mainly that there is no significant difference between P. equisetiforme var. graecum and var. peyerinhoffi supporting a taxonomic grouping of these two varieties.

Klíčová slova:

Epidermis – Flowers – Pollen – Stamens – Plant ovary – Stigma – Anthers – Style


1. Brummitt RK. Vascular plant families and genera. Royal Botanic Gardens, Kew; 1992.

2. Heywood VH. Scanning electron microscopy in the study of plant materials. Micron. 1969; 1: 1–14.

3. Budel JM, Farago PV, Duarte MdR, Takeda IJ. Morpho-anatomical study of the cladodes of Homalocladium platycladum (FJ Muell.) LH Bailey (Polygonaceae). Rev Bras Farmacogn. 2007; 17: 39–43.

4. Koochak H, Seyyednejad SM, Motamedi H. Preliminary study on the antibacterial activity of some medicinal plants of Khuzestan (Iran). Asian Pac J Trop Med. 2010; 3: 180–184.

5. Li A. Polygonaceae AL Jussieu. Flora of China. 2003; 5: 277–350.

6. Brandbyge J. Polygonaceae Flowering Plants· Dicotyledons: Springer; 1993. pp. 531–544.

7. Ronse Decraene LP, Akeroyd J. Generic limits in Polygonum and related genera (Polygonaceae) on the basis of floral characters. Bot J Linn Soc. 1988; 98: 321–371.

8. Pottier-Alapetite G. Flore de la Tunisie: Angiospermes-Dicotyledones. 1. Apetales-Dialypetales. Ministère de ĺ Enseignement Supérieur et de la Recherche Scientifique et le Ministère de ĺ Agriculture; 1979.

9. Haraldson K. Anatomy and taxonomy in Polygonaceae subfam. Polygonoideae Meisn. emend. Jaretzky. Acta Universitatis Upsaliensis, Symbolae Botanicae Upsalienses; 1978.

10. Lersten N, Curtis J. Foliar anatomy of Polygonum (Polygonaceae): survey of epidermal and selected internal structures. Plant Syst Evol. 1992; 182: 71–106.

11. Yasmin G, Khan MA, Shaheen N, Hayat MQ. Micromorphological investigation of foliar anatomy of genera Aconogonon and Bistorta of family Polygonaceae. Intl J Agric Biol. 2009; 11: 285–289.

12. Hedberg O. Pollen morphology in the genus Polygonum s.l. and its taxonomic significance. Svensk Botanisk Tidskrift. 1946; 40: 371–404.

13. Hong SP, Hedberg O. Parallel evolution of aperture numbers and arrangement in the genera Koenigia, Persicaria and Aconogonon (Polygonaceae). Grana. 1990; 29: 177–184.

14. Nowicke JW, Skvarla JJ. Pollen morphology and the relationship of the Plumbaginaceae, Polygonaceae, and Primulaceae to the order Centrospermae. Smithsonian Contributions to Botany. 1977; 37: 1–64.

15. Barthlott W. Epidermal and seed surface characters of plants: systematic applicability and some evolutionary aspects. Nord J Bot. 1981; 1: 345–355.

16. Martin AC. Identifying polygonum seeds. The Journal of Wildlife Management. 1954; 18: 514–520.

17. Yurtseva O, Yakovleva N, Ivanova Radkevich T. Heterocarpy in Polygonum aviculare L. s. str. and related species (Polygonum, subsect. Polygonum). Bulletin of Moscow Society of Naturalists. 1999; 104: 13–20.

18. Ronse Decraene LP, Hong SP, Smets E. Systematic significance of fruit morphology and anatomy in tribes Persicarieae and Polygoneae (Polygonaceae). Bot J Linn Soc. 2000; 134: 301–337.

19. Bauer R. Entwicklungsgeschichtliche untersuchungen an Polygonaceenbluten. Flora. 1922; 115: 272–292.

20. Emberger L. 1939. Aperçu général sur la végétation du Maroc: commentaire de la carte phytogéographique du Maroc 1: 1 500 000. H. Huber Berne; 1939.

21. Ronse Decraene LP, Smets E. The floral development of Popowia whitei (Annonaceae). Nord J Bot. 1990; 10: 411–420.

22. Hildebrand F. Die geschlechter-vertheilung bei den pflanzen. Engelmann, Leipzig; 1867.

23. Hassan M, Khan M. Style-stamen dimorphism in Polygonum L. (Polygonaceae). Bangl Jf Bot. 1987; 16: 93–95.

24. Reddy NP, Bahadur B, Kumar PV. Heterostyly in Polygonum chinense L. J Genet. 1977; 63: 79.

25. Chen ML, Zhang XP. Distyly in Polygonum jucundum Meisn.(Polygonaceae). Plant Syst Evol. 2010; 288: 139–148.

26. Lloyd D, Webb C. The evolution of heterostyly. Evolution and function of heterostyly: Springer; 1992. pp. 151–178.

27. Hong SP. The dimorphic heterostyly in Aconogonon campanulatum (Polygonaceae). Plant Syst Evol. 1991; 176: 125–131.

28. Kantachot C, Chantaranothai P. Achene morphology of Polygonum sl (Polygonaceae) in Thailand. Trop Nat History. 2011; 11: 21–28.

29. Chen ML. Floral morphology and breeding system in Polygonum hastato-sagittatum Mak.(Polygonaceae). Flora. 2012; 207: 365–371.

30. Huang LJ, Fu WL, Wang XF. Floral development at multiple spatial scales in Polygonum jucundum (Polygonaceae), a distylous species with broadly open flowers. PloS one. 2014; 9: e102802. doi: 10.1371/journal.pone.0102802 25058669

31. Wang CH, Du W, Wang XF. Reproductive investment in a cleistogamous morph of Polygonum jucundum (Polygonaceae). Plant Syst Evol. 2017; 303: 559–563.

32. Schneider CA, Rasband WS, Eliceiri KW. NIH Image to ImageJ: 25 years of image analysis. Nat Methods. 2012; 9: 671–675. doi: 10.1038/nmeth.2089 22930834

33. Faivre AE. Ontogenetic differences in heterostylous plants and implications for development from a herkogamous ancestor. Evolution. 2000; 54: 847–858. doi: 10.1111/j.0014-3820.2000.tb00085.x 10937258

34. Sampson BJ, Stringer SJ, Marshall DA. Blueberry floral attributes and their effect on the pollination efficiency of an oligolectic bee, Osmia ribifloris Cockerell (Megachilidae: Apoidea). HortScience. 2013; 48: 136–142.

35. De Castro CC, Araujo AC. Distyly and sequential pollinators of Psychotria nuda (Rubiaceae) in the Atlantic rain forest, Brazil. Plant Syst Evol. 2004; 244: 131–139.

36. Koch AK, Silva P, Silva CA. Biologia reprodutiva de Psychotria carthagenensis (Rubiaceae), espécie distílica de fragmento florestal de mata ciliar, Centro-Oeste do Brasil. Rodriguésia-Instituto de Pesquisas Jardim Botânico do Rio de Janeiro. 2010; 61: 551–558.

37. Deng Y, Chen S, Teng N, Chen F, Li F, Song A, Guan Z. Flower morphologic anatomy and embryological characteristics in Chrysanthemum multicaule (Asteraceae). Sci Hortic. 2010; 124: 500–505.

38. Rao BH, Rao PP. Sporogenesis and gametogenesis of some Boraginaceae. Feddes Repertorium. 1992; 103: 35–40.

39. Zulkarnain Z. Embryology of Swainsona formosa (Fabaceae): anther and ovule development. J Bioscience. 2005; 12: 11.

40. Chehregani A, Tanomi N. Ovule ontogenesis and megagametophyte development in Onobrychis schahuensis Bornm.(Fabaceae). Turk J Bot. 2010; 34: 241–248.

41. Zou F, Sheng-Lin C, De-Yi Y, Zhang RQ, Zhang L, Xiong H. Microsporogenesis, megasporogensis and male and female gametophyte development in Feijoa sellowiana (Myrtaceae). Int J Agric Biol. 2016; 18.

42. Zhang Q, Hao Q, Guo X, Liu Q, Sun Y, Liu Q, Wang K. Anther and ovule development in Camellia japonica (Naidong) in relation to winter dormancy: Climatic evolution considerations. Flora. 2017; 233: 127–139.

43. Yang S, Peng H, Liang H. Embryological observation on Camellia yunnanensis var. camellioides with a comparison of embryological features in Camellia. Guihaia, 2002; 22:340–344.

44. Ghimire B, Suh GU, Lee CH, Heo K, Jeong MJ. Embryological studies on Abelia tyaihyoni Nakai (Caprifoliaceae). Flora. 2018; 242: 79–88.

45. Hong SP, Oh IC, De Craene LR. Pollen morphology of the genera Polygonum s. str. and Polygonella (Polygoneae: Polygonaceae). Plant Syst Evol. 2005; 254: 13–30.

46. Yurtseva O, Severova E, Bovina IY. Pollen morphology and taxonomy of Atraphaxis (Polygoneae, Polygonaceae). Plant Syst Evol. 2014; 300: 749–766.

47. Zhang X, Zhou Z. A study on pollen morphology and its phylogeny of'Polygonaceae in China. China Science and Technology University Press; 1998.

48. Yasmin G, Khan MA, Shaheen N, Hayat MQ. Taxonomic significance of leaf epidermal anatomy of selected Persicaria Mill. species of family Polygonaceae from Pakistan. Afric J Biotechnol. 2010; 9: 3759–3768.

49. Nuraliev MS, Sokoloff DD, Oskolski AA. Floral anatomy of Asian Schefflera (Araliaceae, Apiales): Comparing variation of flower groundplan and vascular patterns. Int J Plant Sci. 2011; 172: 735–762.

50. Von Balthazar M, Endress PK. Development of inflorescences and flowers in Buxaceae and the problem of perianth interpretation. Int J Plant Sci. 2002; 163: 847–876.

51. Laubengayer RA. Studies in the anatomy and morphology of the polygonaceous flower. Am J Bot. 1937; 24: 329–343.

52. Tucker SC. Floral ontogeny in Sophoreae (Leguminosae: Papilionoideae). III. Radial symmetry and random petal aestivation in Cadia purpurea. Am J Bot. 2002; 89: 748–757. doi: 10.3732/ajb.89.5.748 21665674

53. Tucker SC. Overlapping organ initiation and common primordia in flowers of Pisum sativum (Leguminosae: Papilionoideae). Am J Bot. 1989; 76: 714–729

54. Sundberg MD. Short communication petal‐stamen initiation in the genus Cyclamen (Primulaceae). Am J Bot. 1982; 69: 1707–1709.

55. Bello MA, Rudall P, González F, Fernández-Alonso JL. Floral morphology and development in Aragoa (Plantaginaceae) and related members of the order Lamiales. Int J Plant Sci. 2004; 165: 723–738.

56. Endress PK. Floral structure and evolution of primitive angiosperms: recent advances. Plant Syst Evol. 1994; 192: 79–97.

57. Lord JM. Flower and fruit. Morphology, ontogeny, phylogeny, function, ecology. New Zeal J Bot. 2011; 49: 141–142.

58. Neubauer BF. The development of the achene of Polygonum pensylvanicum: embryo, endosperm, and pericarp. Am J Bot. 1971; 58: 655–664.

59. Richards J, Koptur S. Floral variation and distyly in Guettarda scabra (Rubiaceae). Am J Bot. 1993; 80: 31–40.

60. Yurtseva O, Kuznetsova O, Mavrodieva M, Mavrodiev E. What is Atraphaxis L.(Polygonaceae, Polygoneae): cryptic taxa and resolved taxonomic complexity instead of the formal lumping and the lack of morphological synapomorphies. PeerJ. 2016; 4: e1977. doi: 10.7717/peerj.1977 27168986

61. Roth I. Fruits of angiosperms. Encyclopedia of plant anatomy. 1977.

62. Costea M, Tardif FJ. The biology of Canadian weeds. 131. Polygonum aviculare L. Can J Plant Sci. 2005; 85: 481–506.

63. Yurtseva O. Ultrasculpture of achene surface in Polygonum section Polygonum (Polygonaceae) in Russia. Nordic J Bot. 2001; 21: 513–528.

64. Le Floch É, Boulos L, Vela E. Catalogue synonymique commenté de la flore de Tunisie. Simpact; 2010.

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