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Intraspecific variation in the pollen exine sculpture of Jaborosa runcinata Lam. (Solanaceae)

Mara C. Tellera ab; Gloria Daners c a CONICET. Laboratorio de Sistemtica y Biologa Evolutiva, Museo de La Plata, Paseo del Bosque s/n, La Plata, Argentina b Laboratorio de Actuopalinologa, Museo Argentino de Ciencias Naturales, Ciudad Autnoma de Buenos Aires, Argentina c Dpto. de Evolucin de Cuencas (Palinologa Aplicada), Facultad de Ciencias, Universidad de la Repblica, Montevideo, Uruguay Online Publication Date: 01 January 2007 To cite this Article: Tellera, Mara C. and Daners, Gloria (2007) 'Intraspecific variation in the pollen exine sculpture of Jaborosa runcinata Lam. (Solanaceae)', Grana, 46:4, 268 - 273 To link to this article: DOI: 10.1080/00173130701647030 URL: http://dx.doi.org/10.1080/00173130701647030

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Intraspecific variation in the pollen exine sculpture of Jaborosa runcinata Lam. (Solanaceae)

A C. TELLERI A1,2 & GLORIA DANERS3 MARI

CONICET. Laboratorio de Sistema tica y Biolog a Evolutiva, Museo de La Plata, Paseo del Bosque s/n, La Plata, a, Museo Argentino de Ciencias Naturales, Ciudad Auto noma de Buenos Aires, Argentina, 2Laboratorio de Actuopalinolog n de Cuencas (Palinolog a Aplicada), Facultad de Ciencias, Universidad de la Repu blica, Argentina, and 3Dpto. de Evolucio Montevideo, Uruguay
1

Abstract The variability of ectexine sculpture in the pollen of Jaborosa runcinata (Solanaceae) was studied with light and scanning electron microscopy. Three main ectexine variants are recognized: incomplete reticulate, a combination of incomplete reticulate and gemmate-granular, and gemmate-granular. A range of transient conditions were also noted between the two extremes. Apertures vary from a basically triporate-aspidate arrangement to pollen grains which appear to have a rudimentary colpus. The sculptural variability of the ectexine, and a tendency towards a colpus, in J. runcinata are interesting as intraspecific variation does not appear to be common within Solanaceae.

Keywords: Incomplete reticulate, granular, gemmate, Ubisch bodies, aspidate apertures, rudimentary colpus

During the preparation of a pollen reference collection for use in the identification of the pollen content of Uruguayan honey, we noticed that the pollen exine of Jaborosa runcinata showed strong sculptural variability. Both granular and reticulate pollen grains were evident. Jaborosa (Solanaceae), originally described as Trechonetes Miers, is a genus of 22 herbaceous species with an extensive distribution ranging from southern Peru, Paraguay, Bolivia, Brazil, and Uruguay to Tierra del Fuego (Barboza & Hunziker, 1987; Barboza & Romanutti, 1999). All species occur in Argentina. The pollen of this genus appears to have been examined first by Erdtman (1952) in a comprehensive pollen morphological study of angiosperm families. He noted that the pollen of Jaborosa caulescens (5J. bergii Hieron. ssp. caulescens) is 3colporate (brevicolpate) and reticulate while the pollen of J. integrifolia Lam. is 35 aperturate (aspidate) tenui-exinous, and scattered with spinuloid or granuloid excrescences. In a taxonomic study

of Jaborosa, Barboza (1986) examined the pollen morphology of 18 of the 22 species that grow in Argentina. Later, Barboza and Hunziker (1987) used both pollen morphology and macromorphological data to divide the genus into two sections: Jaborosa and Lonchestigma. Section Jaborosa comprises: J. runcinata, J. integrifolia, J. odonelliana and J. oxipetala, and is characterized by tri- or tetraporate pollen with a tectate ectexine covered by supratectal processes. The remaining 18 species of Jaborosa are all referred to section Lonchestigma, which has tricolporate pollen, and a semitectatereticulate ectexine without supratectal processes. In an atlas of pollen for north-eastern Argentina, Cabrera and Cuadrado (2001) describe the pollen of J. runcinata from Buenos Aires Province as triporate with reticulate sculpture while the pollen of another collection from Santa Fe Province is granular. The taxonomic usefulness of pollen morphological data in Jaborosa has already been demonstrated (Barboza & Hunziker, 1987), and a range of pollen

Correspondence: Mar a C. Teller a, Laboratorio de Sistema tica y Biolog a Evolutiva, Museo de La Plata, Paseo del Bosque s/n, 1900, La Plata, Argentina. E-mail: telleria@netverk.com.ar (Received 12 October 2006; accepted 29 June 2007) ISSN 0017-3134 print/ISSN 1651-2049 online # 2007 Taylor & Francis DOI: 10.1080/00173130701647030

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characters has previously been described for the genus (Erdtman, 1952; Barboza, 1986; Cabrera & Cuadrado, 2001). Therefore, the intraspecific pollen variation shown in the specimens of J. runcinata, which we collected, has prompted us to document our data. Material and methods The flowering period for Jaborosa runcinata lasts from October to March. During that period four populations were sampled and pollen from mature anthers of at least five specimens in each population was examined; usually only one open flower per plant is found. Voucher specimens from each population have been deposited in the Herbarium of La Plata (LP) and in the Herbarium de la Ca tedra de Bota nica de la Facultad de Qu mica of Uruguay (MVFQ) (see Results section). Ten flowers from two of the four populations were selected for more detailed studies. For light microscopy (LM), pollen grains were acetolyzed following the protocols outlined in Erdtman (1960), and mounted in glycerol jelly. The different sculpture patterns of the ectexine were defined and counted on the slides. Counts were made on ca. 100 pollen grains obtained from flowers of two populations; one in Uruguay: Montevideo (34 53 S and 56 10 W) and the other in Argentina: Miramar (38 15 S and 57 52 W). The pollen measurement data are based on LM observations of 25 grains per specimen, using an Olympus BX40 microscope in Uruguay and a Nikon E2000 in Argentina. LM photomicrographs were taken with a Nikon Coolpix 4500 camera. Although most counts were based on pollen of entire androecia from the seven Argentinean (Miramar) flowers, the percentages of sculpture variability at single anther level and individual pollen sac level were also investigated in one of the three Uruguayan (Montevideo) flowers (Table I). For SEM studies, because acetolysis causes pollen grains to shrink, non-acetolyzed pollen grains were examined. Pollen grains were suspended in 90% ethanol, mounted on stubs, sputter-coated with gold palladium and examined with a JEOL JSM T-100 microscope. Terminology follows Punt et al. (2007). Herbaria abbreviations are according to Holmgren et al. (1990).

Table I. Percentages of the three sculptural variants in the ten flowers from two populations of Jaborosa runcinata. Anthers- Individual IR Populations Flowers whole anther sacs % Montevideo 1 A B C 2 3 Miramar 4 5 6 7 8 9 10 Ai Aii 58 64 68 37 16 15 IR + Ge-Gr Ge-Gr % % 25 27 20 36 33 29 25 10 70 44 17 9 12 27 50 56 100 75 90 100 30 56 100

IR incomplete reticulate; IR + Ge-Gr a combination of incomplete reticulate with gemmate-granular; Ge-Gr gemmategranular. 1(A-C): counting from pollen sacs or an individual anther of the same flower; 2-10: pollen counted unselectively from anthers within the androecium of an individual flower.

& 2B, E), polar outline rounded triangular (Figures 1D, E, F & 2C, D, F), polar length 45 55 mm, equatorial diameter 45 55 mm. Apertures 3-porate (rarely, 4-porate) and aspidate (Figure 1C, E, F). SEM images show some of the grains to have an incomplete reticulate ectexine (Figures 1A, B, 2B, C, D & 3A, B) with a differentiated area above and below each pore, which appears to be a rudimentary colpus (Figure 3A), although this is not clearly observed with LM. Exine: 1.5 2.5 mm thick, a much thicker and more compact aspis is formed around the pores. In addition, usually small (of similar size to the granular structures) Ubisch bodies are occasionally observed in all variants, see for example, Figure 2B (arrow). We recognize three main variants of ectexine sculpture: 1. Incomplete reticulate [IR] (Figures 1A, B, 2B) 2. A combination of incomplete reticulate and gemmate-granular [IR + Ge-Gr] (Figure 2C, D) 3. Gemmate-granular [Ge-Gr] (Figures 1D, E, F, 2E, F & 3C, D). In this variant the sculptural elements may be more or less evenly distributed (Figure 2F) or randomly aggregated (Figures 1D, 2E). The infratectum is columellate in the incomplete reticulate variant and, in the granular variant; the granula are more or less without columellae and in direct contact with the foot layer. In SEM the muri usually show only very short columellae (Figure 3B). Interestingly we noticed that the smaller grains tend to have a thicker exine than larger grains.

Results Pollen morphology Pollen isopolar, radially symmetrical and suboblate to spheroidal in shape: P/E 0.811.00 (Figures 1A, B

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Figure 1. LM of Jaborosa runcinata pollen. A-C. Slightly tilted equatorial view of pollen with incomplete reticulum. (A) High focus. (B) Low focus. (C) Mid focus showing the thin exine. D-F. Polar view of pollen with a gemmate-granular ectexine. (D) High focus. (E) Mid focus. (F) Mid focus showing the aspidate pores. Scale bars 8 mm.

Ectexine sculpture pattern variation We calculated the relative proportions of the three main sculpture variants (Table I) and it was evident that in most flowers both whole anthers and individual pollen sacs showed sculptural variability in the two populations. Nevertheless, in spite of a

general tendency to variability within individuals, pollen from each of the two populations was also observed to have a high percentage of only one of the sculpture variants: incomplete reticulate in the Uruguayan (Montevideo) population, and gemmate-granular or gemmate in the Argentinean (Miramar) population.

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Figure 2. SEM images of the pollen of Jaborosa runcinata. A. General view of within sample variation. B. Equatorial view of pollen with incomplete reticulum, note large Ubisch body (arrow). C, D. Polar views of pollen grains with incomplete reticulate areas interspersed with a few gemmae. E. Pollen grain with gemmate-granular ectexine, equatorial view. F. Pollen grain with granular ectexine, polar view. Scale bars 10 mm.

Discussion There is a notable range of variation in the ectexine sculpture of Jaborosa runcinata pollen in all the collections we studied. As far as we know variability such as this has not been previously recorded in the

pollen of Solanaceae. Studies of the pollen morphology in other species of the family have described pollen with two sculpture types, for example in Markea and Merinthopodium (Persson et al., 1994) but not at the level of individuals. In Solanum there are dioecious species where the male flowers

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Figure 3. Close up SEM images of the pollen of Jaborosa runcinata. A. Equatorial view showing the expanding pore with expanding aperture membrane, and the rudimentary colpus with microgranular surface structures. B. Detail of incomplete reticulum showing very short columellae, in association with the muri, over pollen grain surface C. Equatorial view showing an aspidate pore with aperture membrane in place. D. Detail of gemmate-granular exine. Scale bars 2 mm.

produce viable 3-zonocolpate pollen while the female flowers usually have inaperturate nonfunctional pollen (Knapp et al., 1998). The ectexine sculpture observed in the pollen of Jaborosa runcinata appears to show a continuum of variation ranging from an interrupted or incomplete reticulum, through gemmae and finally a complete loss of tectum in the free or relatively free and randomly distributed granula, of the granular variant. It is interesting to note that, although some Ubisch bodies have been observed in all three variants, as far as we have observed they are more particularly associated with the gemmate-granular variants. Pollen wall ultrastructure has not been examined with transmission electron microscopy (TEM) and it is difficult to elucidate clearly from LM and SEM alone. However, the granular and gemmate exine elements appear to be compact, and closely adhered to the foot layer, and the columellae, in the incomplete reticulate variant, are extremely reduced (Figure 3B).

A modification of the sculptural elements to granula, adjacent to the pores, appears to be indicative of rudimentary colpi. This tends to be particularly associated with the incomplete reticulate exine variant (Figure 3A), whereas pollen with a gemmate and/or granular ectexine is more likely to have a thick aspis surrounding the pore (Figures 1E, F & 3C). These modifications suggest a trend from pollen with a stronger, more complete tectum (reticulate w incomplete reticulate) to a granular variant where the exine compensates for a reduction in general wall strength by the development of a localized protective thickening around the aperture and more densely distributed granular sculptural elements. Pollen ectexine sculpture variability counts for Jaborosa runcinata show that anthers and individual pollen sacs frequently produce grains where either the gemmate-granular or incomplete reticulate variants predominate over the incomplete reticulate/gemmate-granular variant. In combination with the variation in aperture morphology also observed,

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it seems unlikely that a sampling taken from only one or two specimens would be sufficient to fully characterize the pollen morphology of J. runcinata. Furthermore, the fact that there is usually only one flower open at a time for each plant is another reason why our results, based on a wider sampling, have demonstrated variability not previously observed by authors such as, Erdtman (1952), Barboza (1986) and Cabrera and Cuadrado (2001). Conclusions In summary, the pollen of Jaborosa runcinata, not previously observed to be variable within species, has been demonstrated to have both ectexine sculpture and aperture variability. Developmental studies of the pollen of this species might lead to a better understanding of whether these modifications to ectexine sculpture, and to aperture structure occur during the sequences of growth and development of the microspore (Borsch & Wilde, 2000) or whether, at least for exine variability, it might be the result of a genetically programmed phenomenon, related to the exine template, which occurs during the selfassembly process (Gabarayeva & Hemsley, 2006). Specimens examined
Jaborosa runcinata. URUGUAY: Dpt. Montevideo, Daners 4204 (MVFQ); Dpt. Flores, Daners 4305 (MVFQ). Jaborosa runcinata. ARGENTINA: Prov. La Plata, Buenos Aires, Teller a s.n. (LP); Miramar, Teller a s.n. (LP).

Stefan Vinckier who provided valuable comments, which greatly improved the manuscript. This work was supported by Consejo Nacional de Investigaciones Cient ficas y Te cnicas (CONICET). References
Barboza, G. E. (1986). Estudios palinolo gicos en Jaborosa Juss. y Trechonaetes Miers. (Solanaceae). Bol. Acad. Nacl Cie. (Co rdoba), 57, 357376. Barboza, G. E. & Hunziker, A. T. (1987). Estudios sobre Solanaceae. XXV. Revisio n de Jaborosa. Kurtziana, 19, 77153. Barboza, G. E. & Romanutti, A. (1999). Solanum. In F. O. Zuloaga & O. E. Morrone (Eds), Ca talogo de las plantas vasculares de la Repu blica Argentina. III. FabaceaeZygophyllaceae (Dicotyledonae) (pp. 10691071). St. Louis, MO: Mo. Bot. Gard, Monogr. Syst. Bot. 74. Borsch, T. & Wilde, V. (2000). Pollen variability within species, populations, and individuals, with particular reference to Nelumbo. In M. M. Harley, C. M. Morton & S. Blackmore (Eds), Pollen and spores: Morphology and biology (pp. 285299). Kew: R. Bot. Gard. Cabrera, M. M. & Cuadrado, G. (2001). Morfolog a pol nica de las Solanaceae del Nordeste Argentino, Subfamilia Solanoideae Tribu: Solanae, Datureae, Jaborosae y Lycieae. In S. M. Pire, L. M. Anzo tegui & G. A. Cuadrado (Eds), Flora pol nica del Nordeste Argentino, 2 (pp. 125136). Corrientes: EUDENE. Erdtman, G. (1952). Pollen morphology and plant taxonomy. Angiosperms. Stockholm: Almqvist & Wiksell. Erdtman, G. (1960). The acetolysis method, a revised description. Sv. Bot. Tidskr., 54, 561564. Gabarayeva, N. & Hemsley, A. R. (2006). Merging concept: The role of self-assembly in the development of pollen wall structure. Rev. Palaeobot. Palynol., 138, 121139. Holmgren, P. K., Keuken, W. & Schofield, E. K. (1990). Index Herbariorum P. 1. The herbaria of the World. 8th ed. Bronx, N.Y.: IAPT & NYBG, Regnum Veg., 120. Knapp, S., Persson, V. & Blackmore, S. (1998). Pollen morphology and functional dioecy in Solanum (Solanaceae). Pl. Syst. Evol., 210, 113139. Persson, V., Knapp, S. & Blackmore, S. (1994). Pollen morphology and systematics of tribe Juanulloeae A. T. Hunziker (Solanaceae). Rev. Palaeobot. Palynol., 83, 130. Punt, W., Hoen, P. P., Blackmore, S., Nilsson, S. & Le Thomas, A. (2007). Glossary of pollen and spore terminology (2nd ed. rev.) Utrecht: Rev. Palaeobot. Palynol., 143, 181.

Acknowledgements We thank John Skvarla for critical reading of the manuscript, and Viviana Barreda for helping with the LM photographs and the Curators of the Herbarium of La Plata and the Herbarium of Ca tedra de Bota nica de la Facultad de Qu mica of Uruguay for their assistance. We would also like to thank our reviewers Andrew E. Pozhidaev and