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I. Primexine development in Passiflora racemosa Brot.: overlooked aspects of development

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Abstract

Developmental stages during the tetrad period were examined in detail by transmission electron microscopy with an emphasis on substructure. Our purpose was to find out whether the sequence of sporoderm developmental events provides additional evidence for our recent hypothesis on the underlying cause of exine ontogeny as a sequence of self-assembling micellar mesophases initiated by genomically given physicochemical parameters. Osmiophilic globules encrusting the surface of postmeiotic microspores and tapetal cells are temporary prepattern units which come first. The second prepattern structures are highly ordered bundles of microfilaments and microtubules which determine the position of microspore surface invaginations and clusters of the glycocalyx inside them. The first glycocalyx units are microgranules which during the middle tetrad stage rearrange into radially oriented rod-like units. The latter form lens-like clusters of the glycocalyx-1, located inside the invaginations. These clusters predestine the position of the future luminae in the exine reticulum. The second glycocalyx layer is laid down as a continuous layer over the whole microspore surface and has similar substructure, that is radial rods. Glycocalyx-2 is a framework for procolumellae which appear at the late tetrad stage. Therefore, the sequence of substructural units in the primexine is: globules, microgranules, rod-like units, and layers of radially oriented rods tightly packed in the periplasmic space. This sequence corresponds to the first three mesophases of self-assembling micelles: spherical micelles, cylindrical micelles, and layers of hexagonally packed cylindrical micelles (middle mesophase). We observed the same sequence in other species during primexine development, and the findings of this study provide new evidence for our hypothesis.

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Abbreviations

AL:

Anther loculus

C:

Callose

Co:

Columella

D:

Dictyosome

ER:

Endoplasmic reticulum

G:

Glycocalyx

G1:

Glycocalyx 1

G2:

Glycocalyx 2

GV:

Golgi vesicles

LG:

Lipid globule

mc, MC:

Microspore cytoplasm

Mi:

Mitochondrion

MSP:

Microspore surface pockets

N:

Nucleus

P:

Plastid

PM:

Plasma membrane

PS:

Periplasmic space

RER:

Rough endoplasmic reticulum

RV:

Reticulum vesicles

SER:

Smooth endoplasmic reticulum

Ta:

Tapetum

V:

Vacuole

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Acknowledgments

This work was supported by grant RFBR No. 11-04-00462a to Nina Gabarayeva. We thank our engineer Peter Tzinman for assistance with the Hitachi H-600 transmission electron microscope. We particularly thank Bruce Sampson for correcting the English text. This study on Passiflora exine development was reported at IPC XIII 2012, at the symposium SS10 “Exine development and pattern formation, unifying ultrastructural and genetic approaches” (28 August, Tokyo, Japan) which was dedicated to the memory of the outstanding palynologist John R. Rowley. We cordially thank the co-organizers of the symposium Steve Blackmore and Michael Hesse who did much in preparing this meeting. We heard seven interesting talks, including the introduction by Steve Blackmore, reviewing the past, present and future of exine development studies, and his concluding remarks. The studies, reported at the symposium showed that ideas of Rowley continue to inspire new investigations of different aspects. John Rowley did not set up official “schools of thought”, but such a school has appeared spontaneously, without his special effort.

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Gabarayeva, N., Grigorjeva, V. & Kosenko, Y. I. Primexine development in Passiflora racemosa Brot.: overlooked aspects of development. Plant Syst Evol 299, 1013–1035 (2013). https://doi.org/10.1007/s00606-013-0757-2

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