The structure of individual variability of pollen morphological features of the genus Physalis L. (Solanaceae) and methods of its interpretation (genesis or morphoses)

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Abstract

The morphology of pollen grains in samples of 42 species of the genus Physalis L. was studied using light and electron scanning microscopes. Pollen is morphologically fairly uniform. Although the intrageneric subgroups (subgenera and sections) are heterogeneous in terms of pollen morphology, it is not possible to draw boundaries between them, since the same variants of characters are found in different subgenera and sections. SEM-detected sculpture details are good additional features for the characterization of some species. The pollen of P. alkekengi, sometimes isolated as a monotypic subgenus Physalis, or a separate genus, slightly differs from all other Physalis in micro-verrucate-echinate sculpture. In samples of 19 species (44% of those studied), various rare pollen forms deviating from the typical ones in the number and arrangement of apertures were found. On the example of morphological features of pollen of the genus Physalis and other unrelated taxa, the properties of the complete individual variability of the gametophyte generation are described as an extreme model (having maximum completeness with minimal complexity) for studying the properties of variability. A non-typological description and interpretation of the observed properties of individual morphological variability is given. The own properties of variability (continuity, transitive ordering, parallelism) are described and are not fully provided for in the typological and phylogenetic approach, in which the genus-species discrete-hierarchical ordering of biodiversity is considered universal. The described properties of variability show that during asexual reproduction in the gametophyte generation of pollen (ontologically complete and genetically uniform; haploid, unicellular, spherical living bodies), the ancestral type of organization is not inherited (the typical form is taxon-specific, physiologically normal, adaptive, functional), but with a certain frequency in the generation (in the absence of genetic variability), morphosis arise (form rebirth – non-inherited, but regularly occurring variability), which add up to the series of metamorphosis. In the full generation of the genealogical line, there is not the genesis of the form (generation, the emergence of one form from another), but morphosis (rebirth of the form, change in generic characteristics, archetype, body scheme, without changing the species characteristics). Individual variability can be described as metamorphosis, the fluidity of an individual form, according to J. W. Goethe (complete sets of morphoses, forming a continuous and successive morphological series of living bodies between various typical forms), or the indeterminacy of discrete forms in a continuous series of variability.

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A. E. Pozhidaev

Komarov Botanical Institute, RAS

Author for correspondence.
Email: pae62@mail.ru
Russian Federation, Prof. Popov Str., 2, St. Petersburg, 197376

V. V. Grigoryeva

Komarov Botanical Institute, RAS

Email: mikhailov_val@mail.ru
Russian Federation, Prof. Popov Str., 2, St. Petersburg, 197376

A. N. Semenov

Komarov Botanical Institute, RAS

Email: grigorieva@binran.ru
Russian Federation, Prof. Popov Str., 2, St. Petersburg, 197376

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2. Fig. 1. Morphology of typical pollen of representatives of the genus Physalis (CM, SEM): a–d – P. alkekengi; d’ – P. glabripes; ef–k – P. lobata; l–p – P. lanceifolia; p' – P. acuminata. a, c, f, h, l, n – view from the pole; b, d, g, i, m, o – view from the equator; d, d’, j, p, p’ – surface sculpture (SEM). a, b, f, g, l, m – CM; c–d, d’, h–j, n–p, p’ – SEM. The upper and lower focus are shown for each pollen grain (CM). Scale bars, µm: a–g, f–i, l–o – 10; d, d’, j, p, p’ – 1.

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3. Fig. 2. Morphology of typical pollen grains of the genus Physalis (CM, SEM): a–d – P. pruinosa; f–j – P. ixocarpa; k–p – P. angustifolia. a, c, f, h, l, n – view from the pole; b, d, g, i, m, o – view from the equator; d, j, p – surface sculpture (SEM). a, b, f, g, l, m – CM; c–d, h–j, n–p – SEM. The upper and lower focus are shown for each pollen grain (CM). Scale bars, µm: a–g, f–i, l–o – 10; d, j, p – 1.

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4. Fig. 3. Morphology of typical pollen grains of the genus Physalis (CM, SEM): a–d – P. parviflora; d’ – P. nicandroides; е–к – P. missouriensis; л–п – P. lagascae. a, c, e, h, l, n – view from the pole; b, d, g, i, m, o – view from the equator; d, d’, j, p – surface sculpture (SEM). a, b, e, g, l, m – CM; c–d, d’, h–j, n–p – SEM. The upper and lower focus (CM) are shown for each pollen grain. Scale bars, µm: a–g, e–i, l–o – 10; d, d’, j, p – 1.

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5. Fig. 4. Deviating forms of the arrangement of pollen apertures of the genus Physalis (CM): a, e, g – P. gracilis; b–d – P. virginiana; h, k, p, t – P. adulterina; and – P. subintegra; l, n, y – P. macrophysa; m – P. angustifola; o – P. angulata; p – P. parviflora; c – P. сinerascens; f – P. mollis. a – form A; b – form B; c, d – form G; d – form D (typical T); e – form B; g – form Ж (solid-aperture, tennis ball-shape); h, i, l – form Ж (separate aperture W-shape); m – form Z; n, p, p – form I; o – form E; c – form L; t – form M; y, f – deflecting 3-colporate pollen grain with oblique arrangement of endoapertures. B, V, D, Z, L – intermediate forms of aperture arrangement. For letter designations of forms, see Fig. 5. The upper and lower focus (CM) are shown for each pollen grain. Scale bars are 10 µm.

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6. Fig. 5. Schematic representation of the diversity of pollen forms of the genus Physalis and the periodic structure of ordering of the diversity of pollen forms of angiosperms. a – Schematic representation of the diversity of pollen forms of the genus Physalis (A–M – diversity of pollen forms of the first period, T – typical pollen grains); reduced images at the bottom to the right of forms G, D, Zh, I, K and M show the same forms in a different projection. b – Distribution of the occurrence of pollen forms by the number of species of the genus Physalis (the height of the columns and the numbers for each form in the diagram indicate the number of species in which the indicated form was found).

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7. Fig. 6. Parallelism of typical and deviant pollen forms of some taxa of angiosperms. a–c – Krameria secundiflora DC.; g – K. grayi Rose; d–g – K. cistoides Hook.; h – K. ramosissima S. Wats; and – K. erecta Willd. ex Shultes; j – K. parviflora Benth (Krameriaceae); l – Acer maybei Maxim. (Sapindaceae); m – Eupomatia laurina R. Br. (Eupomatiaceae); n – Scutallaria dubia Taliev et Schirj. (Lamiaceae); o – Acer davidii Franch. (Sapindaceae); n – Monstera deliciosa Liebm. (Araceae); p – Pedicularis peduncularis Popov (Scrophulariaceae); c – Alsodeia echinocarpa Korth. (Violaceae); t, y – Acanthosyris falcata Griseb. (Santalaceae); f – Euphorbia leptocaula Boiss.; x – E. saravschanica Regel (Euphorbiaceae); c – Eugenia luzonensis Snerr. (Myrtaceae). a–i, m, p, r – typical pollen grains (marked with the letter T in the upper right corner); k, l, n, o, s–c – atypical pollen forms. a–c – 3-colpate form D; g–zh – acolpate-3-ore form D; z, i – acolpate, endoaperture of form A; k – acolpate, endoaperture of form Zh (tennis ball form); z, i, m–r – form A; k, l – form Zh (tennis ball form); s–c – form E (s, t – right variant; x, c – left variant). For letter designations of forms, see Fig. 5. a–c – typical pollen grains of p. Krameria (Krameriaceae) with exoapertures; g–f – without exoapertures. a, g, e, h, c, t, h, c – SM; b, c, e, f, g–r, u, f – SEM. Upper and lower focus (SM) are shown for each pollen grain. Scale bars – 10 µm.

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