Dr. Robert Heiling Library, S.m.s.medical College, Jaipur
Pollen And Spore Morphology Plant Taxonomy
Front matter
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Almquist & Tiksell/Gebers Förlag AB
STOCKHOLM 1957
IN SWEUCIV
PRINTED IN SEDEN BY
Almqvist & Wiksells
BOKTRYCKERI AKTIEBOLAG
UPPSALA 1957
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6
с
d
Juniperus contunis, part of an acctolyzed megaspore membrane at different
adjustnients of the mit roscope from high (a) te lon (d) , 1000)
POLLEN AND SPORE
MORPHOLOGY/
PLANT TAXONOMY
GYMNOSPERMAE, PTERIDOPHYTA, BRYOPHYTA
(ILLUSTRATIONS)
(An Introduction to Palynology. II)
BY
>
G. ERDTMAN
Irish fróntiếprece, 5 plates and 265 illustrations
ALMQVIST & WIKSELL/ STOCKHOLM
TJIE RONALD PRESS COMPANY / NET YORK
Almquist & Wiksell Gebers Förlag AB
STOCK IOLM 1957
IN SWEUCI
PAINTED IX SREDEX BY
Almerisi & Tiksells
BOKTRYCKERI AKTIFBOLAG
UPPSALA 1957
PREFACE The second part of "An Introduction to Palynology" deals, as was originally planned, with pollen and spore morphology in the Gymnospermae, Pteridophyta, and Bryophyta. Although the illustrations and most parts of the text were ready for printing in the autumn of 1956, the completion of the text had to be somewhat postponed on account of my absence abroad for several months It was accordingly decided to publish the second part in two instalments thus, the present volume (Vol. II) consists mainly of illustrations, whereas the following volume (Vol. III) will provide the related text, Most of the illustrations are palynograms drawn after the author's originals by Anna-Lisa Nilsson They depict, in a standardized manner, a pollen grain or spore in polar and/or lateral view, together with its sporoderm stratification and other details. It should emphasized, however, that the sporoderm stratification details merely hint at the great array of patterns and layers which exist. Pending thorough morphological investigations, the proper interpretation of these layers etc. is still, in many cases, largely conjectural. Besides the palynograms, some photomicrographs and electron micrographs of very thin sections through spore walls are included Stimulated hy my eminent friend, Dr R, W Kolbe, the microscopist and diatom specialist, I tentatively took up—in the late nineteen-forties--electron microscopy as an aid in sporoderm research. These studies have been continued, with better technique and greater success, by Dr. Barbro Afzelius, to whom I am indebted for the electron micrographs here reproduced (with the exception of those of Pl. IV and V, which were put at my disposal by Dr. D. von Wettstein) and for a chapter (pp. 125-131) on the application of new microscopical techniques in palynological research. Another chapter (pp. 135–147), by J. Radwan Praglowski, deals with the technique of making very thin sections from spore material embedded in plastic. As shown by Fig. 1 (P.2), this technique can also be applied in the study of very old (Palaeozoic etc.) spores. In conformity with Vol. I, the present volume has been prepared under the auspices of the Swedish Natural Science Research Council. Help and support of various kinds received from other sources will be duly acknowledged in Vol. III. Palynological Laboratory, Stockholm - Bromma, July 1957. G.E.
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CONTENTS
3
INTRODUCTION
On the exine morphology of the saccate pollen grains in recent
gymnosperms
3
5
ILLUSTRATIONS
5
Gymnospermae (Abies—Viddringtonia)
Pteridophyta (Acrostichum-Xiphopteris)
45
Bryophyta (Acroschisma-Voiha)
99
SUPPLEMENT
125
On new methods in physical cell research and their application
in studies of pollen grains and spores. By B. M AFZELIUS. 125
On the cutting of ultra-thin sections. By J. RADWAN PRACLOWSKI 135
INDEX
148
Fig 1. Selosisporites sp. (material supplied by Dr Gerhard Kremp), section (thickness about 05 p) through part of the sclerine at high (Icft) and low (riglit) adjustment of the microscope x 1000.
INTRODUCTION ON THE EXINE MORPHOLOGY OF THE SACCATE POLLEN GRAINS IN RECENT GYMNOSPERMS The saccate pollen grains in recent gymnosperms are heteropolar, bilateral or radiosymmetric (sometimes slightly asymmetric). They consist of a body (corpus) and a varying number of airsacks or bladders (s accı). The aperture is distal, and should often perhaps more appropriately be referred to as a tenuitas (i e., a thin aperturoid area functioning as an aperture and gradually merging into the surrounding evine). It has earlier, as a rule, been described as a sulcus or a sulcoid groove. The surface of the corpus of a pollen grain with n bladders can be divided into the following areas. n saccale areas, forming the floor of the saccı, nmesosaccale areas (mesosaccia), 1.e. areas between the sacci and in the same latitude as these, and finally two a posaccale areas (a posaccia), one at the distal pole, and the other, usually much larger than the former, in the proximal face of the grain with the proximal pole in its centre. With respect to the thickness of the exine of the corpus certain pollen types (cf. e.g. Figs. 53 and 57) exhibit two distinct exine areas a proximal, crassi-exinous (referred to as cap, cappa), and a distal, tenui-exinous (referred to as cappula). The non-saccale esine of the cappa consists of comparatively thick sexine and thin nexine. The outer, ectosexinous part of the sexine is usually thin, tegulloid, and as a rule connected to the nexine-except within the saccl--by baculoid, densely spaced endosexinous elements. The sacci are separated from the interior, non-exinous parts of the corpus, by saccale nexine. Their outer wall consists of thin ectosexine which is often perforated (shown in electron micrographs—not published by Erdtman and Thorsson in 1950). The small holes (micropuncta) are usually difficult to observe through an ordinary light microscope. In the majority of the Tsuga species the ectosexine of the saccı (as well as that of the corpus, cf. Fig 73) is studded with spinules or small spines. Attached to the inner surface of the outer wall of the sacci are endo-sexinous elements protruding into the lumen of the bladders (in Pherosphaera filzgeraldu stray endosexinous rods are also found on the saccale nexine). These elements are more widely spaced than those of the body. Branched or unbranched, single or combined in different ways, they tend to produce an array of patterns which are difficult to draw and hard to describe. Microtomc sections (cf particularly Afzelius in Grana palynologica, 1:2, 1936) make these subtle details of pollen construction casier to observe and saler to interpret. Ncar the proximal root of the sacci arc often found slight, scxinous ridges or frill-like projections (proximal crests, cristac proximales, also referred to as cristae marginales) varying in appearance in different species. At the distal root of the sacci, where these merge into the distal aposaccium, the characteristic pattern of the bladders comes abruptly to an end. The height of the corpus coincides with the polar axis (i.e. the perpendicular line connecting the potes); the breadth is identical with its maximum horizontal extension in grains in cquatorial longitudinal view (marginal crests extending beyond the general surface of the corpus not included), and the depth (in bilateral grains) is equal to the transverse ("non-sacciferous") diameter of the corpus. It is often prcierable and, at the same time, easier to calculate the inner dimensions of the corpus. The height of a saccus is the shortest distance from the highest point of the saccus (or from a line drawn through this point parallel to the saccale nexine) to the underlying nexine of the corpus. Its breadth is equivalent to the "tangential" diameter of the saccus in pollen grains in polar view. In radiosymmetric grains the breadth can also be measured in pollen grains seen in equatorial view; bilateral grains must be in a transverse equatorial position iſ the breadth shall be measured. In microscope slides this, lowcver, is seldom the case. Its depth-—ın bilateral grains is equal to the maximum diameter of the saccus in grains in equatorial longitudinal view. In radiosymmetric grains the depth is calculated is a similar way. Height, breadth, and depth of corpus and sacci are illustrated in Figs. 2, 3, 13, 23, 27, 43, 53, 57, 58, and 62. As shown by Afzelius the inner part of the nexine (the endonexine) in Cedrus is laminated (Fig. 12, p. 11). In acetolyzed polien grains of this genus, and of Abies, etc., it can often be secn, even by means of an ordinary microscope, that the nexine consists of two distinct layers, which often split apart as a result of the chemical treatment. So striking is this feature that it seems extraordinary that it has not been mentioned until now. The morphology of bisaccate pollen grains has been dealt with by numerous botanists, among whom was Strasburger, who believed that the floor of the sacci was formed by intine. This opinion has often been echoed right up to recent years, although Strasburger himself soon corrected his mistake. In conclusion, it ought to be mentioned that, according to Ciguriaeva, the thin ends (with sexine and nexine slightly separate from each other) of the pollen grains in Ephedra and JVelwitschia may be interpreted as the remainder of true sacci.
ILLUSTRATIONS
GYMNOSPERMAE
Frontispiece; Pl. I (facing p. 12), Pl. II (facing p. 20); Figs. 2–73.
MEGASPORES
Cupressaceae: Frontispiece; Figs. 36- Pinaceae. Figs. 44 (p. 28), 52 (p 32),
41 (pp. 23-25).
63 d (p.40).
Podocarpaceae. Fig. 19 (p. 14).
POLLEN GRAINS
Araucariaceae: Figs. 5 (p. 8), 7 (p. 9). Gnetaceae. Figs. 34, 35 (p 22).
Cephalotaxaceae: Pl. 1 (Fig. 2); Fig. Pinaceae: Figs. 2 (p 6), 12, 13 (p. 11),
14 (p. 12).
13 (p. 27), 45 (p. 28), 53 (p 33),
Cupressaceae. Figs. 4(p. 8), 11(p 10), 55-58 (pp. 31–36), 62 (p. 39), 63
29 (p. 18), 42 (p. 26), 49 (p.30), 54 a-c (p.40), 73 (p. 41)
(p. 31), 71 (p. 43).
Taxaceae. Figs. 6 (p. 8), 9 (p. 9), 50
Cycadaceae Figs. 10 (p. 10), 17 (p. (p. 31), 70 (p 42)
13), 18 (p. 13), 28 (p. 18), 31 B Taxodiaceae. Figs. 8 (p. 9), 15 (p 12),
(p. 20), 16 (p. 29), 67 (p. 42). 16 (p. 12), 33 (p. 21), 47 (p. 29),
Ephedraceae: Pl. II; Figs. 30 A, B 65, 66 (p. 11), 68, 69 (p. 12), 72
(p 19), 31 A (p. 20).
(p. 13).
Ginkgoaceae: Fig. 32 (p. 21). Welwitschiaceae Fig. 30 C (p. 19).
ABIES:-
B
001
-
in
A
Fig. 2 A, Abies nephrolepis; a, lateral longitudinal view, b, lateral transverse view; c, polar view (distal face). X 650. B, A. magnifica, lateral longitudinal view. x 650. C, A. mariesit, saccus pattern (about x 2500) 1: distal pole, 4 proximal pole Corpus height 1-4, breadth 5-5, depth 3-3. Sacci: height 9-8, breadth 10-10, depth 11-11. Total grain: height 4-12 (A: a, b), breadth 13-13, depth 3-5-3 (A: b), 3-2-1-2-3 (A: c) Eleren of the inserted photomicrographs show the pattern of the outer surface of the sacci at different foci (the uppermost photomicrograph in each saccus shows the pattern at high adjustment of the microscope, the lowermost the same at low adjustment) Two of the photomicrographs between numerals 11 and 8 in Aa and B show a part of the saccus in optical cross-section. The remaining four photomicrographs exhibit the pattern of the corpus Those in a b are phase contrast pictures. (From Erdtman in Svensk bot Tidskr. 1954) ACMOPYLE:- 13 13 W2106 10 Fig. 3. dcmopyle pancheri. For explanation of numerais see Fig. 2.
8
ACTINOSTROBUS:
0
Fig. 4. Actinostrobus acuminatus, surface and optical cross-section. x 1000.
AGATHIS:--
A
B
ь с
Fig. 5. A, Agathis microstachya, crine stratification x 2000. B, A. ovala,
erine stratification (a, * 2000) and LO-patterns (1-d).
AMENTOTAXUS:-
or
pr
Fig. 6 Amentolarus argotaenia; di, distal pole; pr, proximal pole. From left
to right: three still loosely united pollen grains (tenuitas distal; 250); exine
stratification (X 2000); lateral view, surface and optical cross-section (x1000);
LO-patterns; pollen grains in polar view (x 250).
ARAVCARIA: b Fig 7. Araucaria montana (exinc stratification). 1, transition from tenuitas (left) to the thickwallet rest of the grain (right) Layers interpreted in the same way as in b. -- b antic, exine stratification outside tcnuitas (alternative interpretations); e, shows a thin "nesine" (possibly consisting of two layers) overlain with a thick sexine" exhibiting three layers: an "endosesinous faintly-marked layer (consisting of baculoid rods?) merging into a thick, + granular "ectosexine" ("tegilum"), beset with piloid "suprategillar" processes. According to the interpretation given in b, thin "setine" "pila") covers a thick "nexine" (the rest of the exinc). - d, LO-patterns ("pila"). * 2000, ARCEUTHOS: Fig. 42 B, p. 26. ATIROTAXIS:-- Fig. 8. Athrotaris cupressades, From left to right: erine stratification (2000); lateral view of opened grain (* 1000); LO.patterns AUSTROTAXUS:-- Fig. 9. dustrolarus spicali. Vain figure: surface and optical cross-section (* 1000). Delails. exine stratification (* 2000) and 1.0-patterns.
BOVENIA: EI Fig. 10 Bowenia spectabilis. Details marked 3 x cnlarged 3000 times. The main figure shows the distal facc (surface and optical cross-section), x 1000. CALLITRIS:-- D Fig. 11 Callitris balansae var. alpina. Main figure surface and optical crosssection (* 1000) Details (from left to right)lateral view of opened grain (* 250); exine stratification (* 2000); LO-patterns. CEDRUS:- C. atlantica (Pl I (facing p 12); Fig. 12 a, b, C. deodara (Fig. 13).
5 Fis 12. Cedrus atlantica 2, section through part of an acetolyzed pollen wall; from top to bottom' ectoscunt, endosexinous rods, et tonerine (homogeneous), cndonexine (laminated, the blach horizontal line indicates 1 * 22.000); , enconerine (from another section, magnification greater than in a). ENG B M. Afzchius 164 (221 13 D Fig. 13. Cedrus deodara. For coplanation of numerals see Fig 2, p. 7. 2-563817 O Erdıman CEPITALOTAXUS:- A B Fig. 14. &, Cephalolaxus druprcear; a, pollen grain with eraginated tenuitas (lateral vicw; < 1000); b), pollen grain with ini aginated tenuitas (distal face, * 1000) 1, C. nana, exine stratification (* 2000). — The drawing, for this figure were ready before the electron micrograph, Pl. I, Fig. 2. was taken. The white counterpait ("enconexine") to the lamellar layer in the EMG is clearly seen Superimposesi on the "ondonerine" is a black layer ("ecto. nexine") corresponding, it would scem, to the granular part of the “nexine" shown in the EMG. Also with regard to serine details there is an obvious similarity between Fig. 11 and the EMG, MI I, l'ig. 2. CHIAMAECYPARIS- Megaspore membrane: Chamaecyparis lawsoniana, see Fig. 36, p. 23. l'olien grains: Chamaecyparis pisfera, see Fig. 42 A, p. 26. CRYPTOMERIA Fig 15 Cryptomeria japonicn From left to right pollen grain in lateral view, surface (* 250); pollen grain (ilistal face invaginated) in lateral vies, surface and optical section (* 1000), crine stratification (* 2000). > CUNNINGHAMIA: Fig. 16. Cunninghamia lanceolata From leit to right distal face (upper detailfigure; <250), LO-patterns (lower detailfigure); lateral view, surface and optical section (ilistal face invaginated; x 1000); cxine stratification (x 2000).
Fig 1 Cedrus allantron, section through part of an acetolyzed pollen wall The nexine has been split off ſiom the sevne at the point indicated by the arrow The endonexine is laminated 18,000 The black horizontal line indicates 1 (CMG, from Afzelius, in Grana palynologica, 1 2, 1956 ) l'ig 2. Cephalotaras naina, cune stratification 12,000. The black hori. zontal line indicates 1 # (LMG, from Afzelius, in Grana palynologica, 1. 2, 1BAY
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IJ CUPRESSUS: Megasporc membrane: Cupressus arizonica, sec lig 37, p 23, C whitleyann, sec Fig 38, p. 24. CYCAS (sce also Fig. 31 B, p 20) – Fig 17. Cycas revoluta; outline of transverse median section (upper detailfigure), proximal face (lower detail-figure). Fig 18. Cycas revoluta, sections through part of acetolyzed pollen wallscf tangential. + obliquc section (left), + radial section (right, 34,000) The black horizontal line indicates 1 CMG B M. Afzelius DACRYDIUM: Megaspore membrane: D. cupressinum (Fig. 19). Pollen grains D. araucarioides (Fig. 20 B), D. bidwillii (Fig. 21). D. ela. lum (Fig. 20 A), D. falciforine (Fig 22), D). fonkii (Fig. 23) D franklinii (Fig 24), D guillaumini (Figs. 25 and 26), D. laroides (Fig. 27).
2012 Fig. 19. Dacrydium cupressinum; megaspore membrane (* 2000) and LOpatterns of sanie. B Fig. 20. A, Dacrydium elatun; exine stratification in proximal aposaçciuni (* 2000) and pollen grain in lateral, longitudinal view (surface; nevine contours indicated by broken lines; X 1000). B, D. araucarioides, pollen grain in lateral, longitudinal view (surface; distal face invaginated as indicated by the broken line just below the centre of the figure; * 1000), a, proximal face (* 250); b, distal facc (* 230).
Fig 21. Dacrydium bidwillii, distal face (left) and pollen grain in lateral, longitudinal view (surface, optical section, and LO-patterns) * 1000 acrydum falciforme, a, lateral view, b, proximal face b 16 8 13 A 13 13 Fig. 23. Dacrydium fonkit, upper figure: outline of pollen grain in lateral, longitudinal view ( 1000) Lower figure: distal face, surface and optical section ( - 1000). For ctplanation of numerals sce Fig. 2. p. 7. The curved brohen linc in the extreme right part of the lower figure indicates the ap. proxiniate delimitation of a concavity in the distal lace of the saccus. 10 b Fig. 24. Dacrydium franklinii; a, lateral, longitudinal view; , proximal face; C-1, $ deviating grains (c distal, d proximal face); C, LO-patterns; f, evinc stratification, proximal face (* 2000; 2-1 * 1000).
17 Fig 25. Dacrydium guillauminu From left to right. lateral view (* 250). distal face (* 1000), LO-patterns at high and successively lower focus Tug 26 Dacryduim guillauminu; oblique view (slightly > x 1000).
13 113 10 Fig. 27. Dacrydium taroides. Upper figure: pollen grain in lateral, longitudinal view, surface and section. Lower figure: proximal face, surſace (left) and optical secfion (right; ~ 1000). Lower left-hand detail-figure: outline of pollen grain in lateral, transverse view(x250;ouler contour of the corpus marked by the broken line) For explanation of numerals see Fig. 2, p 7. DIOON -- ♡ 20 Fig. 28. Dioon edule; upper lenhand detail figure: median sagittal section; lower left-hand detailfigure: frontal section; upper righthand detail-figure: transierse sec tion (all x 1000); lower righthand detail-figures: evine stratification (* 1000) and LO-patterns. DISELMA: Fig. 29. Diselna archeri; surface and optical section (* 1000); erine stratification (* 2000); LOpatterns.
19 EPHEDRA [Fig. 30 A, B, Fig. 31 A, PI. II (facing p. 20)], WELWITSCHIA (Fig. 30 C): – 0 b A B C Fig. 30. A, Ephedra antisyphıltıca, uppermost detail-figure: section through the exine along line x-y in the main figure (* 2000), b), LO-patterns; the detail-figure (X 2000) between this figure and the main figure (* 1000) shows a section through the evine at one of the short ends. B, E. equisetina; cf distal face (* 1000), surface and optical section, c, young grain with sulcoid aperture. C, Welwitschia mirabilis, cf. lateral, transverse view (x1000), exine stratification (2000), and cf distal face (* 1000), uppermost detailfigure: LO-analysis of part of the exine surface (to be read from left to right); as the two central details show there is a very low and narrow ridge (hright in the second, dark in the third detail from the left) at the bottom of the valleys (stippled in the extreme left, white in the extreme right detail) separating the hollow serine ridges (white in the extreme left, stippled in the extreme right detail) Fig. 31. A (main figure) Ephedra monosperma; section through an acetolyzed pollen wall, * 19,000. B (lower right-hand corner). Cycas revoluta, section through part of an acetolyzed pollen wall. * 35,000. The black horizontal lines indicate 1 4. (ENG; from Afzelius Grana palynologica, 1:2, 1936.)
PII Ephedra monosperma, section through part of the inner laminated laver suf an acetolyzed pollen wall (the inner part of the pollen grain is to the len, the black horizontal line indicales 11) $13,000 (CVG; from Afzelius in Grana palynologica, 1 2, 1956.)
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21 FITZROYA: see Fig. 42 G, H, p. 26. GINKGO:- 0000 Fig. 32. Ginkgo biloba; extreme left detail-figure pollen grain in lateral, longitudinal view with an outline of the accompanying inedian transversc section Second detail-figure from the left pollen grain slightly tilted with the entrance to the concavity which lodges the tenutas shown to the left. The remaining figures exhibit the distal face of an unexpanded and an expanded grain. The lower detail figure of the latter gives the outline of a pollen grain in lateral transverse view (owing to an omission a full line uniting the upper extremities of the figure is not shown) * 1000 GLYPTOSTROBUS:--- Fig. 33. Glyplostrobus pensils. From left to right: opened grain (* 250), lateral view, surface and optical section (* 1000). crine stratification (* 2000), LO-patterns Uppermost detail-figure: tenuitas (optical section; Y 2000).
22 GNETUNI: А B с Fig 34. A, Gnetum venosum. – B, G. africanum. - C, G. montanum. (From Erdiman in Bot. Notiser 1954.) Fig. 35. Gnetum montanum, section through part of an acetylyzed. very compressed pollen grain. To the extreme right is a spinule. The black horizontal line (indicating 1 q) is drawn within the lumen of the grain x 21.000 EMG R M. Afzelius. JUNIPERUS, ARCEUTHOS, CHAMAECYPARIS, CUPRESSUS, FITZROYA, LIBOCEDRUS, WIDDRINGTONIA Megaspore membranes: Chamaecyparis lawsoniana (Fig. 36, p. 23). Cu. pressus arizonica (Fig. 37, p 23). C. whilleyana (Fig. 38. p 24), Juniperus communis (Frontispicce and Fig. 39, p. 24), J. sabina (Fig. 41, p 25). Pollen grains. Arcenthos drupacea (Fig 42 B. P 26), Chamaecyparis pisiferat (Fig 42 A), Fitzroya cupressoides (Fig 42G, H), Juniperus californica (Fig. A2 D), J prostrata (Fig. 42 E), Libocedrus decurrens (Fig. 42 F), Widdring. tonia cupressoides (Fig. 42 C).
Fig. 36. Chamaecyparis lawsoniana, megaspore inembrane (surface). phase contrast. x 4000. (From von Lurzer in Grana palynologica, 1:2, 1956.) Fig. 37. Cupressus arizonica; megaspore membrane (surface). phase contrast. X 4000. (From von Lürzer in Grana palynologica, 1: 2. 1936 ) Fig 38 Cupressus whilleyana, megaspore membrane (surface), phase contrast. * 1000 (From von Lurzer in Grana palynologica, 1.2, 1936) Fig 39 Juniperus communis; megaspore membrane (optical section). * 1000 (For iO-patterns of the membrane surface, see Frontispiece.)
Fig. 40. Juniperus subuna, megaspore membrane (optical section), phase contrast. X 4000. (From von Lurzer in Grana palynologica, 1.2, 1956.) Fig. 41. Juniperus sabina, megaspore membrane (surface), phase contrast x 4000 (From yon Lurzer in Grana palynologica, 1:2, 1936) B A ♡ G 50. F G H Fig. 42. Cupressaceae. A, Chamaecyparis pisifera B, Arcenthos «supacea C. Widdringtonın cupressoides D. Juniperus californica. E, J. prostrata. F Libocedrus decurrens, G, H, Fitzroya cupressoides. The main figures (* 1000) exhibit grains in lateral view with (in A-C, and F-H) the distal pole at the top. An aperturoid spot is faintly marked in the upper left-hand quadrant in D. The detail figures show the cxine stratification enlarged 2000 times. The detail-figures between the main figure and the LO-patterns in E show an irregular grain (upper detail-figure) and an opened grain (both x 250), H is a Filcroya pollen grain of normal type whereas in G à deviating, "subsaccate" grain is shown - Pollen grains of other cupres saceous plants are shown in Figs. 4 (Actinostrobus). 11 (Callitris),29 (Diseima), 49 (Neocallitropsis), 51 (Pilgerodendron), and 71 (Thujopsis).
KETELEERIA:
8
13
1
10
13
10
Fig. 43, Keteleeria davidiana; lateral, longitudinal view and distal face
(* 1000). For explanation of numerals see rig. 2, p 7. (From Erdman in
Svensk bot. Tidskr. 1931)
3-365617 G. Erdiman
LARIX Megasporo membrane: Larir occidenfolis (Fig. 44). Pollen grains: Larix decidua f. polonica (Fig. 45 1), 1.. gmelini var, japonica (Fig. 45 A), L. occidentalis (Fig. 45 C). Fig 44 Larix occidentalis, acctolyzed megaspore wall, optical section (above) and surface (below). X 1000. А B Fig. 45. A, Larir gmelini var. japonia; cxine stratification and Lo-patterns. B, L. decidua f. polonica; exine stratification anul "Lo-patterns", C, 1.. occidentalis, lateral view, surface and optical section (* 1000).
29 LIBOCEDRUS: see Fig. 42 F, p. 26. MACROZAMIA:- Die a Fig. 46. Macrozamia spiralıs; a, transverse, equatorial view (optical crosssection); b, distal face, c, evne stratification at amb at right angles to central part of the sulcoid tenuitas, d, pattern caused by endosexinous rods (phase contrast); e-g, morphological variants a, b x 1000, < X 2000, e-8 * 250. METASEQUOIA:- : Fig. 47. Metasequoia glyptostroboides, lateral view (* 1000) and exine stratification (* 2000).
MICROCACHRYS (Fig. 48 A), MICROSTROBUS (Fig. 48 B):-
O
00
А
B
Fig. 48 A, Vicrocachrys tetragona; a, oblique lateral view (outline: the
sexine of the proximal face in the upper right half part of the figure should
be thinner), b, proximal face (* 1000)B, Microstrobus (Pherosphaera)
fitzgeraldii; c, lateral view (outline); <d, distal face; e, proximal face outline;
* 1000).
MICROSTROBUS: see Fig. 48 B.
NEOCALLITROPSIS:-
Fig. 19. N'eocallıtropsis araucarioides. Main
figue surface and optical cross-section
(* 1000) Details: opened and unopened
grains (* 250); exine stratification (x 2000)
31
NOTHOTAXUS.--
Fig 50 Notholarus chienu From left to right:
lateral view, surface and optical section (* 1000);
eune stratification (2000), LO-patterns
PHEROSPHAERA, see Fig 48 B, p 30.
PHYLLOCLADUS:
g
di
di
90
x di
Fig. 51. Phyllocladus protraclus, a, lateral, longitudinal view of a pollen
grain (* 306) with the proximal face covered by a "saccus" (z), possibly
an artificial feature resulting from the loosening of the sexine from the
ncxine, b, g, lateral (longitudinal and transverse) view (surface), c, f, same
as b and g, in optical section; d, distal face (* di - distal pole). e, proximal
face (* pr = proximal pole) – 5-6 X 1000.
PICEA:
Megaspore membranc: Picea abies (Fig. 52).
Pollen grains: Picea jezoensis (Fig. 53).
-
b
Fig 52. Picea abies: part of an acetolyzed megaspore membrane at high (a)
and low (b) focus
33
13
4
-
HT?
(511
121
Fig. 53. Picea jezoensis; lateral, longitudinal view and distal face (surface and
optical section, x 1000); for explanation of numerals sec Fig. 2, p. 7.
(From Erdtman in Svensk bot. Tidshr. 1931.)
Ut
PILGERODENDRON:
O
Fig. 54 Pilgerodendron immferum. From left to right two pollen grains (x 250);
distal face (surfaec and aplıcal section, * 1000), exinc stratification (2000),
LO-palferns
PINUS:--
P canariensis (Fig 55), P. excelsa (F26. 57), P. mngo (Fig. 56), P. pence
(Fig 55), P pinen (Fig 55), P thunbergii (Fig. 58)
D
00
760
Fig. 55 Left, Pinus peuce (suhg. Haploxylon). Centre, P. canariensis (subg
Diploxylon, sect. Sula). Right, P pineri (suhe. Diploxylon, sect Pinea)
X 250.
Fig. 56. Pinus mugo; section through saccus and, in the lower right-hand
corner, part of the exine in the proximal facc. About x 1000.
35
RELVA
13
13
Bin
13
D
10
Fig. 57. Pinus eicelsa, lateral, longitudinal view and distal face (surface and
section; * 1000) For explanation of numerals see Fig 2, p. 7. (From
Erdtman in Svensk bot. Tidsbr. 1931.)
(7)
(12)
131
13
13
15
5
EX
12
3 12
13
12
,13
(2)
10
Fig. 58. Pinus thunbergii; lateral, longitudinal view and distal face (surface
and section. * 1000) For explanation of numerals sce Fig. 2. p. 7. (From
Erdiman in Svensk bot. Tidskr. 1954 )
PODOCARPUS:--
P. alpinus var. caespitosus (Fig. 59 C), P. angustifolius var. wrighlü (FA
59 A), P.blumei (Fig G0 B), P. coriaceus (Fig. 59 B), P. dacrydioides (Fig.
60 A), P. minor (Fig. 61 C), P. nagi (Fig. 61 D), P. nubigenus (Fig. 61 B),
P. wallichianus (Fig. 61 A).
37 A B o Fig. 59. A, Podocarpus angustifolius var wrightu, proximal face B, P. coriaceus; distal face. C, P. alpinus var. caespitosus, proximal face y 250. A B b Fig. 60. A, Podocarpus dacrydiordes, a, equatorial view (* 250); b, saccus pattern at high focus; c, pattern of distal face at high (upper detail figure) and low focus; above the main figure (x1000) is an optical section through the evine at the amb (* 2000). — B, P. blumei; saccus paltern and optical section through the outer wall of the saccus.
"A
A
O
B
с
D
Fig. 61. A, Podocarpus wallichianus; lateral, longitudinal view (contour only;
x 250), distal face (* 1000) 1, P. nubigenus. From left to right. proximal
face (* 250); lateral, longitudinal view (1000): LO-patterns (proximal
face) C, P. minor; proximal face (1000) D, P. nagi; lateral, longitudinal
view (* 1000).
39 PSEUDOLARIX: 12 8 13 13 11 3 10 13 13 Fig 62. Pseudolanır amabilis, lateral, longitudinal view and distal face (surface and optical section; x 1000). For explanation of numerals scc Fig. 2, p. 7. (From Erdtınan in Svensk bot. Tidskr. 1931.)
40 PSEUDOTSUGA:- Megaspore mcnibrane: Pseudotsija laxifolin (Fig. 63 d). Pollen grains: Pseudotsuga tarifolia (Fig. 63 a-c). :-1 a d b Fig. 63. Pseudolsuga taxifolia, a, distal pole (with tenuitas); lv, proximal pole; c. part of the microspore exine (optical cross-section, * 2000); <, part of megaspore exine (optical cross-section, * 1900). The upper left-hand detailfigure shows the exine of an opened pollen grain (* 250), the upper right: hand detail-figure a young pollen grain with markings similar to a "tetrad scar". SAXEGOTHAEA: O. Fig. 61. Saregothaea conspicua; lateral view, surface and optical cross-section (x 1000).
SCIADOPITYS: Fig 65. Sciadopitys verticillata, prosimal face (* 1000), evine stratifi (* 2000), and LO-patterns Upper left-hand detail. lateral view pole upwards; * 250) Lower left-hand detail optical section (distal upwards, X 250) SEQUOIA:- 4, B 10# Fig. 66. A, Sequoia (Sequoiadendron) gigantea, lateral view, surface and optical cross-sections (à, 1000, A, * 2000). B, S. sempervirens; lateral view (surface and section; * 1000). 101 101 SEQUOIADENDRON: see Fig. 66 A.
STANGERIA:
a
bo
Fig. 67. Slangeria paradora; distal face (surface and optical section; x 1000):
a lateral view (x250), b, distal face of a grain more broad than long (x 250).
TAIWANIA:-
Fig. 68 Taiwania cryptomerioides. From left to right: cxine stratification
(* 2000), lateral view (* 1000), LO-patterns.
TAXODIUM:-
Fig 69 Tarodum mucronatuin From left to right: erine stratification (x2000);
lateral view (* 1000), LO-patterns
TAXUS:-
O
Fig. 70. Taxus baccata. From left to right: distal face (* 250); pros
face (* 250), lateral view (surface and optical section; ~ 1000); LO-patt
a, cxine stratification (* 2000).
43 THUJOPSIS: 441 Fig. 71. Thujopsis dolabrata, surface (x1000) and exine stratification (2000). TORREYA:- 3x O. g Fig. 72. a, Torreya californica; optical section (* 1000) and exinc stratilication (* 3000) b, t. nucifera, lateral view (surface and optical section; x 1000) and LO-patterns. TSUGA: I. chinensis (Fig. 73 m, o), T diversifolia (Fig 73 g, h), T. dumosa (Fig. 73 ), T forrestii (Fig 73 a-1). T. paltoniana (Fig 73 n), T. sieboldii (Fig. 73 i.)). T yunnanensis (Fig. 73 1). 4- $6.3617 0. Erdiman
OO
བ་ པི་
b
Ал.
с
k
Em A
m
AAN
n
Fig. 73. a-f, Tsuga forrestri; a, distal face (outline; * 250); b, proximal face
(outline of an aberrant grain, * 250), c-e, LO-patterns (c: four spinules (L),
d: the same spinulcs () and upper parts of the undulating tegillum (L),
c: the fleck near the centre (L) is a small infrategillar hollow, also vaguely
shown in d (0)]. f, tegillum. - Left column (6, 1, k, m, n): outline of exine
stratification, proximal face (* 2000), &. T. diversifolia; i, 7 sieboldii:
k, T. dumosa; m, T. chinensis; n, T. palloniana - Right column (h, j, 1, , 0)
tegillum of the puſly frill (* 2000); h, T. diversifolia; j. T. sieboldii; ), T.
yunnanensis, f, T forrestii; o, T. chinensis.
WELWITSCHIA: see Fig. 30 C, p. 19.
WIDDRINGTONIA: see Fig. 42 C, p. 26.
PTERIDOPHYTA Pl. III (facing p 91); Figs 74-191, MEGASPORES Isoetaceae: Figs. 123-125 (pp 67,68) Pilulariaceae: Fig. 162 d, p. 87 Selaginellaceae PI III; Figs. 171– 173 (pp. 90, 91). ISOSPORES ANI) MICROSPORES ? 79). Families in accordance with Engler's Syllabus, 12. Ed , Vol. 1, 1954): Angiopteridaceae: Fig. 144 A-C (p Matoniaceae. Fig. 159 (p. 86), Ophioglossaceae: Figs. 89, 90 (pp 52, Azollaceae. Fig. 85 (p. 50) 53). Christenseniaceae: Fig. 141D (p 79) Osmundaceae Fig 155 (p. 81). Cyatheaceae: Figs. 78 (p. 47), 96 (r Parkeriaceae Figs.89,90 (pp 52,53) 56), 112 (p. 62). Pilulariaccae. Fig. 162 a-c (p. 87). Danaeaceae: Fig. 144 F (p 79) f'lagiogyriaceae: Fig. 163 (p. 88). Dicksoniaceae: Figs. 101 (p 58), Polypodiaceae. Figs. 74–77, 81-81, 169 B (p. 90), 186 (p. 95). 86-88, 91-95, 97-101, 103-107, Dipteridaceae: Fig. 103 (p 58). 109-111, 115-117, 122, 128-132, Equisetaceae: Fig. 106 (p 60). 135, 146, 147, 149, 151, 152, 151, Gleicheniaceae. Fig. 114 (p. 63). 156, 158, 161, 165, 167-169, 183- Hymenophyllaceae. Fig 188 (p 96) 186, 189-191 Hymenophyllopsidaceae. Figs. 119, Protocyatheaceae Figs. 79 (p. 17), 120 (pp. 65, 66). 113 (p. 71), 131 (p. 72) Isoetaceae: Figs. 126, 127 (p. 68). Psilotacene: Fig. 166 (p. 89) Loxsomaceae Fig. 136 (p. 73). Salviniaceae: Fig. 170 (p 90). Lycopodiaceae: Pl. III; Figs. 138 Schizaeaceae: Fig. 143 (p. 78). 142 (pp. 75-77), 161 (p 87). Selaginellaceae: Figs. 174-182 (pp Marattiaceae: Fig. 141 E (p. 79). 91-91). Marsileaceae: Fig. 145 (p 80). Tmesipteridaceae: Fig. 187 (p 96) 16 ACROSTICHUM: l'ig. 74. dcrostichum aureum f. hastacfolitun. Scicrine stratification ( * 2000). a, perine; b), cf. sexine; c, cf, ncxine. ACTINIOPTERIS:- с Fig. 75. Actinopteris australis. Spore in Jateral view (left, * 250); proximal face, surface (a) and optical section (b +c; in b the contours of the verrucae in the distal face are marked, the faint nich in the nexine is quite accidental; x 1000) ADIANTOPSIS: that (?) Fig. 76. Adiantopsis chlorophylla. From left to right: spore in lateral view (* 250); proximal face (* 1000; surface and optical section), LO-patterns ADIANTUM: O rug 77 Adiantum reniforme Provimal fare, surface (left) and optical section (right, x 1000). ALSOPHILA:--- in no dd, b a Fig 78. Alsophila glabra, a, sclerine stratification (2000), , sporc, lateral view (* 250); c, distal face (1000). Surface (left) and optical section (right), d, and le, LO-patterns AMPHIDESMIUM; Tig 70. Amphidesinium blechnoides. Distal face (* 1000), surface (len) and optical section (right). ܕܬܐܟ݂ܗ
ANARTHROPTERIS:
(0
)
Fig. 80. Anarthropteris dictyopteris from left to right' proximal face (* 250),
exinc stratification (with a query, * 2000); spore in lateral view (* 1000),
LO-patterns
ANEMIA: see Fig. 143 G-I, p. 78.
ANGIOPTERIS: see Fig. 144 A, p. 79.
ARCHANGIOPTERIS: see Fig. 141 C, p. 79.
ASPLENIOPSIS:--
Fig. 81. Aspleniopsis decipiens; a and c, deviating spores; b, normal spore
(proximal face; * 1000) Upper right-hand details: lateral view (proximal
face up: * 250); exine stratification (* 2000)
ASPLENIUM Fig. 82. Asplenium monanthes From left to right LO-patterns, sclerine stratification (* 2000), pore in latei al view (* 1000). Fig 83. Asplenium hemonitis From left to right: sclerine stratification (* 2000), distal face (* 1000); LO-patterns. ATHYRIUM: aby Fig. 81. Athyrium filix-femina. From left to right: proximal face (* 250); sclerinc stratification (* 2000), spore in lateral view; LO-patterns.
50 AZOLLA: kan ? Fig. 85. Azolla filiculoides, microsporcs, From above to below: evine stratification (* 2000); proximal face (* 1000, spore surrounded by perine); spore in lateral view (* 1000). BLECHNUM: Fig. 86. Blechnun palmiforme, part of the sclerine (optical section, x 2001 Attached to the inner surface of the exine is a peculiar (abnormal) proce leading into the interior of the spore.
51
BOLBITIS:
Fig. 87. Bolbitis turrialbae. From left to right: proximal face (* 250), sclerine
stratification (* 2000); lateral, longitudinal view (1000)
BOTRYCHIUM: a note on the spores in B simplex is found in the caption of
Fig. 153, p. 83.
BRAINEA:
Fig 88. Brainea insignis. From left to right: lateral, longitudinal view (* 250):
selerine stratification (x 2000); proximal face (* 1000); deviating sporc
(* 250).
CERATOPTERIS:-
Chwa
A
Fig. 89. Ceralopteris siliquosa, proximal face (* 1000): upper detail figures
(from left to right)distal face (x 250), sclerıne stratification (* 2000):
lateral view (* 250) and LO-patterns,
53 b Fig. 90. Ceratopteris sılıquosa Sections through acetolyzed sclerine. The sclerine, as shown in h, consists of a thin perine (the outermost granular layer) and a thick exinc faintly subdivided into cf sexine (showing + parallel ridges separated by U-shaped valleys) and cf. nexine (of equal thickness throughout) The upper figure (a) intimates that the sexine has a fine (granular) structure. 700 CETERACH: ь с a 0 Fig 91. Ceterach officinarum. From left to right: proximal face (250); selerine stratification (2000): lateral, longitudinal view (* 1000); LOpatterns.
54 CHEIROPLEURIA:- b C Fig. 92. Cheiropleuria bicuspis; 2, proximal face (* 1000); b, sclerine stratification (* 2000); c, distal lace (* 1000). CHRISTENSENIA: see Fig. 144 D, p. 79. CIBOTIUM:-- b Fig. 93. Cibotium barometz, proximal face (* 1000). a, lateral view (* 250); b, distal face (* 250). 1 and 2, respectively, are identical.
Fig. 95. Cryptogramma brunoniana. From left to ris).f. lateral virw (27.
distal face (* 1000), exine stratification (2010).
55
CNEMIDARIA:
Fig. 91. Cnemidaria speciosa From left to right: LO-palterns, diutat fare
(* 1000) and sclerine stratification (* 2000),
CRYPTOGRAMMA:
56
CYATHEA (see also HEMITELIA):-
a
b
C
e
Fig. 96. Cyathea vestila a, sclerine stratification (* 2000); }), LO-patterns;
c, distal face (aperture wrongly marked by broken instead of dotted lines);
(I, laesura (open), e, lateral view (* 230).
CYCLOPHORUS: see PYRROSIA, p. 89.
CYRTOMIUM: see PHANEROPHLEBIA, P. 85.
CYSTODIUM: see Fig. 169 B, P.
90.
CYSTOPTERIS:
Fig. 97. Cyslopteris fragilis. From left to right: scicrine stratification (* 2000);
lateral, longitudinal view (* 1000); proximal ſacc (* 250) and LO-patterns
Fig. 98. Cystopteris fragilis dicheana from left to right proximal face
(optical section, * 250), sclerıne statification (* 2000); lateral, longitudinal
view (* 1000), LO-patterns.
DANAEA: see Fig. 144 F, p. 79.
DAVALLIA:
Fig. 99 Davallia canariensis Proximal face (* 250); lateral, longitudinal
View (* 1000), and exine stratification (* 2000).
DENNSTAEDTIA -
Fig. 100 Dennstaedlı punchilobula, From left to right. selerine stratification
(* 2000), proximal face, surface and section (* 1000); analysis of laesura.
DICKSONIA: Fig. 101. Dicksonia youngiae From left to right lateral view (* 250), distal (ui) and proximal (pr, surface and optical section) face; LO-patterris. DIPLAZIUM: Fig 102. Diplazium proliferum. From left to right: proximal face (* 250), sclerine stratification (x 2000); lateral, longitudinal view (* 1000), LOpatterns. DIPTERIS: Fig. 103. Dipteris chinensis From left to right: lateral, longitudinal view (* 250); distal face (* 1000) and exine stratification (* 2000); LO-patterns.
59 DRYMOGLOSSUM:- b B с C Fig. 104. A. Drymoglossum heterophyllum, lateral, longitudinal view (* 1000), a, proximal face ( x 250); b and C, sclerine stratification (> 2000); C, LOpatterns B, D carnosum, selerine stratification (x 2000) DRYNARIA: e Fig 105. Drynaria fortunei. From left to right: proximal face (x 250); scicrine stratification (* 2000), lateral, longitudinal view (* 1000), L.O. patterns 5-565617 G Erdeman
DRYOPTERIS:-
Fig. 106. Dryopteris filix-mas.
From left to right: proximal
facc (optical section, * 250);
sclerine stratification (2000),
cquatorial, longitudinal siew
(* 1000).
ELAPHOGLOSSUM:
T->
Fig. 107. Elaphoglosstim vieillardui. From left to right: proximal face (optical
section, * 250). lateral, longitudinal view (* 1000), sclerine stratification
(* 2000)
EQUISETUM:
b
7
Fig. 108. Equisetum giganteum. a, part
d of scierine of an acetolyzed spore, op
ucal cross-section (* 1000); amplec
tators ("elaters", "hapters") dissolved,
b, sclerine stratification (x 2000); C,
opened spore (x 250). d. part of an
abnormal sporc tetrad (one large, three
small spores; 250).
a
61 GRAMITIS:- Fig 109 Grammitis deplanchei. From left to right lateral view (* 250), proximal face (* 1000), LO-patterns GYMNOGRAMME: Fig. 110. Gymnogramme (Eriosorus) congesta From left to right distal face (* 1000), lateral view (* 250), cf perıne fragments (optical section, x 2000), proximal face (1000) and LO-patterns. HEMIONITIS Fig 111. Hemionitis arifolia From left to right: lateral view (* 250); sclerine stratification (2000). proximal ſace (* 1000) HIEMITELIA: Fig. 112. Hemitelia maxonii. From left to right: lateral view (* 250), proximal face (surface and optical cross-section, 1000); sclerine stratification (* 2000). From Erdiman in Potonić, Svensk bot. Tidskr. 1954. Fig. 113. Hemitelia setosa. From left to right: Jateral view (x 250); distal face, surface and optical cross-section (* 1000); sclerine stratification (* 2000).
63 HICRIOPTERIS (Fig. 114 C), PLATYZOMA (Fig. 114 D), STICHERUS (Fig. 114 B), STROMATOPTERIS (Fig. 114 A):-- C; A 104 Fig 114 Gleicheniaceae. A, Stromatopteris moniliformis, sclerine stratification. B, Sticherus penniger, sclerine stratification C, Hicriopteris laevissima; Cr, proximal face; C,, distal face, Cgpart of laesura (optical cross-section), C, LO-patterns of same D, Platyzoma microphyllum, sclerine stratification. A, B, D x 800, C, and C21000, C, X 4600. HISTIOPTERIS: Fig 115 Hislopteris incisa; proximal facc (* 250) and lateral, longitudinal VICW (optical cross-section and surface, X 1000).
HOLTTUMIELLA:-
Fig. 116 Holtlumiella flabellivenım. Three spores of a tetrad still slightly
adhering x 700
HUMATA:--
Fig. 117. Humala gaimardiana From left to right: latcral, longitudinal view
(* 250), proximal face (* 1000); sclerine stratification (x 2000).
HYMENOGLOSSUM: see Fig. 188 C, p. 96.
65
HYMENOLEPIS:-
Fig 118. Hymenolepis spicata from left to right lateral, longitudinal view
(* 250); distal face (x 1000), a-c, exine stratification showing the successive
increase in thickness (* 2000), LO-patterns.
HYMENOPHYLLOPSIS (Figs. 119, 120).
العون .
Fig 119. Ilymenophyllopsis asplenioides, proximal face, surface and optical
cross-section (* 1000) In some spores the sclerinc stratification appears
as indicated at a, and in other spores as indicated at b,
Fig. 120. Hymenophyllopsis dejecta; distal face (surface and optical crosssection). * 1000. HYMENOPHYLLUM (sce also Fig. 188 C, p. 96);- A 00 0 ارد Fig. 121, A, Hymenophyllum recurrunt; latera) view (* 250) and proximal face, surface and optical cross-section (* 1000). B, H. peltatum, exine stratífication (* 2000).
67 HYPOLEPIS: mur PARTE M A B с Fig. 122. Sclerine stratification in Hypolepis A, H repens B, H. lenufolia C, H. rugosula. Da, H. distans (2000) LO-patterns at lugh (left) and low (right) focus in H distans are shown in D b. ISOETES- Megaspores. Isoetes durieui (Fig 123), 1 echinosporum (Figs 124, 125) Microspores: Isoetes adspersa (Fig 126), 1 baetica (Fig 127). Fig. 123, Isoeles durieui, part of megaspore wall (section). * 1000. 1-25 Fig. 124. Isoeles echinosporum, part of megasporc wall (optical section). The inner contour of the silicified perine is faintly scen left and right of the cf. scrinous outgrowth in the centre of the figure. The "serine" is underlain by thinner "nesine x 2000, Fig 125. Isočles echinosporum, acetolyzed megaspore embedded in glycerine jelly (* 125). In some places the spiny perinc is faintly shown. Fig 126 Isoēles adspersa; sclerine stratification and LO-patterns. A thin cf. perine with hollow, tapering processes, open at top (OL), is underlain by an evine consisting of a tegillate and baculate sexine (LO- bacula) and nevine. 00 C Fig. 127. Isoëtes baelica, a, lateral, longitudinal view (surface); l, lateral, longitudinal view (optical section); c, lateral transverse view (optical section); c, lateral transverse view (optical section); «, polar view (proximal face; sculptine not included), e, exine from a young spore with serine still adhering to the netine; 4000 (and' x 1000).
69
ISOLOMA:
Fig 128. Isoloma divergens From left to right proximal face (* 250), lateral,
longitudinal view (1000), evine stratification
JAMESONIA:
Fig 129. Jamesonia imbricata, proximal face, surface and optical section
(* 1000)
LASTREA:
Fig 130 Lastrea thclypleris. I'rom left to right proximal face (x 250
sclerine stratification (* 2000); lateral, longitudinal view (* 1000): 1
patterns.
LEPTOLEPIA
C
e
b
Fig. 131 Leptolepia norac-Zelandiae, proximal face (* 1000). – a, equatorial
view (* 250), i), proximal face of monolete spore (* 250), c, sclerine
stratification showing perinc (densely dotted) on the outside of the exine;
d. proximal pole at various focuses from high (above) to low, c, general
surface of spore at different focuses.
- N.B. Exceptionally, the "scrrucae"
in the main figure are shown at low focus (i e, dark), and not, as otherwise
in this book, at high focus (bright).
LEPTOPTERIS: sec Fig. 155 G, p. 86.
LINDSAEA:
Fig. 132. Lindsaea orbiculata from left to right: lateral view and proximal
face (both 250); distal face (* 1000); LO-patterns.
71
LOPHOSORIA (Figs. 133, 134):
Fig. 133 Lophosoria quadripinnala (Puerto Rico, Sinfenis 1333, er herb.
Paris); lateral view (surface and section) and proximal face (* 1000) At
top, from left to right distal face (* 250), LO-patterns, selerine stratification
(* 2000)
0 Fig. 134 Lophosoria quadripinnata. Upper figures (Weberbauer 1333, herb. Berlin), from left to right proximal face (* 250), lateral view (* 1000), sclerine stratification (* 2000). Lower figures. proximal face of a Lophosoria spore found in a slide made from Serpyllopsis caespitosa var densifolia (collected by C Shottsberg in the Juan Fernandez Islands August 27th, 1908); a, sclerinc stratification (* 1000); b), lateral view (* 250), c, distal face (x 230).
73
LORINSERIA:-
e
Fig. 135. Lorinseria areolata From left to right: proximal face (* 250);
lateral, longitudinal view (1000), exine stratification (perine not con.
sidered, 2000)
LOXSOMA:
Fig. 136. Loxsoma cunninghami. From left to right: lateral view (* 230):
sclerine stratification (x 2000); proximal face (* 1000): LO-patterns.
LOXSOMOPSIS: Fig 137 Lorsomopsis lehmannir. Lateral view (230), distal face (* 1000), and Lo-patterns At top sclerine stratification (* 2000). LYCOPODIUM:- L. claratum (Pl. III (facing p 94), Figs. 138 B, 142), L. densum (Fig. 139), L. diaphanum (Fig 138 A), L. drummondi (Fig 140). L. insulare Pg 141) Fig. 138. A, Lycopodium diaphanum; a + b), distal face (a. at high, b, at low adjustment of the microscope; the different foci correspond to levels a, and b, in Fig. A: d; c, proximal face (main part of figure: surface view at high adjustment; upper right-hand part: section through the erine; d and e outline of exine stratification at distal pole, c at the equator) – B, L. clavatum: detail corresponding to that in . diaphanum shown in 1'ig A: e. — * 2000 (A: a-c), x 4000 (A: d and e, B), x 500 (lower left-hand detail figure=spore of L. diaphanum in lateral view, distal face up). - From Erdtinan in Afzelius, Erdtman and Sjostrand, Svensk hot. Tidskr. 1954.
15
Fog. 138.
6-565617 a Erdman
Fig. 139. Lycopodium densum; a, distal, b), proximal face (both * 1000); lower right detail-figure: part of evine surface at the transition between the distal and the proximal face. L b Fig. 140. Lycopodium drummondii; a, distal, b, proximal face (* 1000); c, part of laesura at different adjustments of the microscope from high to low. Fig. 141. Lycopodium anulare; distal face ( x 1000) and Lo-patterns.
* ET Тя Fig 142 Lycopodium clavatum, section through an acetolyzed spore wall (x 125,000) cxhibiting the fine, granular-lamellar, structure of the outer part of the cxine The «lıstinct lamellae shown in this figure are multiples of ſine lamellac (thickness 50-60 A). Cach of these consists of a single layer of granules CVG B M Arzelus (From Grana palynologica, 1: 2, 1936.)
78
LYGODIUM (Fig. 143 A-D), ANEMIA (Fig. 143 G-I), MOHRIA (Fig. 143 J),
SCHIZAEA (Fig. 143 E, F):-
A
H
B
.
24
www
m
10
2011
Fig. 143 Schizacaceae, sclerine stratification (* 2000). A, Lygodium circin.
natum. B, L volubile C, L. micans D, L. japonicum. E. Schizaea Numinensis.
F. S. melanesica. G, Anemia phylltidis. H, .. adiantifolia. I. A. anthriscifolia.
J, Nohria caffrorum. (From Erdtman in Svensk bot Tidskr 1951.)
MACROGLENA: see Fig. 188 B, p. 96.
MACROGLOSSUM: see Fig. 144 B,
79.
MARATTIA (Fig. 144 E), ANGIOPTERIS (Fig. 141 A), ARCHANGIOPTENIS
(Fig. 144 C), CHRISTENSENIA (Fig. 144 D), DANAEA (Fig. 144 F), MACRO.
GLOSSUM (Fig. 144 B);-
79 A B Cogo TOTY MY A E D F D. Fig. 144 Marattiales A, Angiopteris longifolia, prorimal face (surface and optical section; 1000). B, Macroglossum alidae, proximal face (surface and optical section; x 1000); C, Archangiopleris henryi; proximal face (surface and optical section; * 1000). D, Christensenia aesculifolia; proximal face (surface and optical section; x 1000); to the left two spores in lateral, trans. verse and lateral, longitudinal view (* 250). E, Jaraltia frarinea; proximal face (surface and optical cross-section; x 1000); to the left two spores in transverse lateral and in longitudinal lateral view (x 250). F, Danaea elliplica; lateral, longitudinal view (* 1000), proximal facc (* 250) and sclerine stratification (* 2000) (From Crdtman in Svensk bot. Tidshr. 1934.)
MARSILEA: 16 Fig. 145. Varsilea aegyptiaca, microsporc. From left to right: lateral view (x 230), proximal face (surface and optical section; * 1000), sclerine stratification (* 2000) and LO-patterns MATTEUCIA A B Fig. 146. A, Matteucia struthiopteris; a, proximal face (x 250); b, lateral, longitudinal view (surface and optical section, X 1000); e, sclerine stralie fication (* 2000), d, LO-paiterns. B, JI. orientalis, LO-patterns. MICROLEPIA:-- Fig. 147. Jficrolepia hırla. From left to right: lateral view (* 250): sclerine stratification (x 2000); proximal face ( 1000); LO-patterns,
MOHRIA: see Fig. 143 J, p. 78
NEGRIPTERIS.
Fig. 148. Negripferis uncana From left to right sclerınc stratification (* 2000);
distal face (surface and optical section, X 1000); LO-patterns.
NEPHROLEPIS
Fug 119 Nephrolepis cordifolia.
From left to right: proximal face
(250), lateral, longitudinal view
(* 1000), sclerine stratification,
(* 2000).
OLEANDRA:---
bile
O
BA
?
Fig. 150 Oleandra neriiformis. From left to right: proximal face (* 250);
lateral view (median optical section; x 1000), selerine stratification (2000)
ONOCLEA: Fig. 151 Onoclea sensibilis from left to right lateral, longitudinal view (* 250); proximal face (surface and optical section; ~ 1000); sclerine stratification, * 2000), LO-patterns ONYCHIUM:-- Fig. 152. Onychium japonicum. From left to right: sclerine stratification (* 2000); proximal face (* 1000); lateral view (* 1000).
83 OPHIOGLOSSUM:- K L M LF AAN -SERI: b e K N 9 0 (pr) Р с Fig. 153. Ophioglossum coriaceum Main figure proximal face (* 1000). surface and optical section, a-d, sections through the erine (2000; a, b, and c are sections along the lines klm, ihno, and op in the main figure (lim corresponds to the letters KLM in fig a ete ) whereas d cthibits a part of the exine with the sexine locally severed from the nexine in Botrychium simplex transitions have been seen from spores with the serine adhering to the nexinc, as in the main figure above, to such where the sexine is almost completely separated from the nerine-due to the chemical treatmentimparting a # pansaccate appearance to the spore); e-8, LO-patterns at high, medium, and low focus. Between d and the main figure is as sporc seen from the side (* 250, di, distal, pr, proximal pole) --Trom Erdtman in Svensk bot. Tidskr. 1954.
ORMOLOMA (Fig. 151 A), ORTHIOPTERIS (Fig. 154 B):
1
B
Fig. 154. A, Ormoloma imrayana; lateral view (* 250), distal facc (*1000),
and sclerine stratification (* 2000). B, Orthiopteris inaequalis; proximal face
(* 1000). exine stratification (* 2000), and-to the left-sporc in lateral
view (* 250) and proximal face of an aberrant spore (* 250).
ORTHIOPTERIS: see Fig. 15+ B.
OSMUNDA (Fig. 155 A-E), LEPTOPTERIS (Fig. 155 G), TODEA (Fig.
155 F):
A
104
PUN
horsrusierung
B
с
E
F
Fig. 155. Osmundaceae, exine stratification (x 2000). A, Osmunda cinna.
momea, part of exine in proximal face (with laesura). B-G, part of exine in
distal face; B, Osmunda regalis (subgen. Euosmunda); C, O. banksiifolin
(subgen. Pleasium), D, O. cinnamomea (suligen. Osmundastrum); E, O:
claytoniana (subgen Osmundastrum); F, Todea barbara; G, Leplopleris
superba,
85 PELLAEA:- Fig. 156 Pellaea viridis From left to ught proximal face (* 250); sclerine stratification (2000), latrial view (1000); LO-patterns PERANEMA:- Fig. 157. Peranema cyatheoides From left to right proximal face (* 250); sclerine stratification ( x 2000); spoie in lateral, longitudinal view (* 1000). LO-patterns (OL-pattern due to small perforations in the perine) PHANEROPHLEBIA: QO0 Fig 158. Phanerophlebia caryolidea From left to right proximal face ( x 250). spore it lateral, longitudinal view (surface and optical section; X 1000); LO-patterns; sclerine stratification (* 2000) 86 PHANEROSORUS: a C Fig 159. Phanerosorus major; a, proximal face, surface and optical sections (* 1000); b, spore in lateral view (* 250), c, monolete spore, proximal face (* 250). The LO-partern in the lower right-hand corner is due to the thin perinous cover PHLEBODIUM: b 250 * 250 * 1000 x 2000 Fig. 100. Phlebodium aureum; a, spore in lateral, longitudinal view, b, proximai face; c, part of proximal face with laesura (broken lines); d, exine stratification, 87 + PHYLLOGLOSSUM: Fig 161. Phylloglossum drummondi, lateral view (* 250) and distal face (* 1000) PILULARIA: Megaspore: Pilularia globulifera (Fig. 162 ) Microspores: Pilularia globulifera (Fig 162 a-c) b 3. 162. Pilularia globulifera. a-f, microspore a, proximal face (surface and optical cross-section; x 1000); 1), laleral view (surface and ophical cross-section, * 1000): C, LO-patterns; d, megaspore, sclerine stratification (* 2000, cf. also Grana palynologica, 1:2, 1936, PI. I, Fig. 13, facing p. 126)
88
PLAGIOGYRIA:
Fig. 163. Plagiogyria henryi.
From left to right: lateral view
(*250); proximalface (250);
distal face (x 250); sclcrine
stratification (* 2000).
PLATYCERIUM:
0
Fig 164. Plalycerium madagascariense. From left to right: proximal face
(* 230), lateral view (surface and optical cross-section; ~ 1000); sclcrinr
stratification (* 2000).
PLATYZOWA; see Fig. 114 D, p. 63.
POLYBOTRYA:
Fig. 165. Polybotrya appendiculata; a, lateral, transverse view (* 250);
h, lateral, longitudinal view (surface and optical section, X1000); c, prori.
mal face (* 250); upper left-hand detail: sclerine stratification (* 2000).
89 PSILOTUM: 6 0 50 100 Fig. 166. Psilotum nudum; a, laesura (optical cross-section; * 2000), b, lateral, longitulinal view (x 250), c, proxinal face (250), d, LO-patterns, e, sclerine stratification 2000) PTERIDIUM Tig 167 Pleridium aquilinunt From Jeſt to right: lateral view ( x 250); sclerine stratification (* 2000); distal face (* 1000) PYRROSIA: Fig 168 Pyrrosia abbreviata, sclerine stratification. (* 2000). SACCOLOMA (Fig. 169 A), CYSTODIUM (Fig. 169 B): IT a Fig. 169 A Saccoloma elegons; 2, sclerine stratification (p, perine?, s, sexine ?, n, nevinc ?, * 2000), b, part of proximal face (pr, pror. imal pole). C, part of distal face (b and C x 1250) Xpr Ь A d Fig. 169 B. Cystodium sorbifoliun, part of distal face (* 1000); d, sclerine stratification, the small rounded excrescences of the thin outermost (perinous ?) stratum are cross-sections of the narrow white ridges exhibited in the main figure (* 2000) B SALVINIA: Fig 170 Salvinia cucullala, microspore: a, lateral view (x 250); b), exine stratification at the transition between the two faces (2000); C, evine stratification, proximal face (* 2000); main figure, proximal face (* 1000) SCHIZAEA: see Fig. 143 E, F, p. 78. SELAGINELLA:-- Megaspores S firmula (Fig 171), S rupestrı (Fig 172), S. selaginoides (PI. III, facing p 94, Fig 173) Microspores: S. apus (Fig 174), S atroviridis (Fig 175), S. biformis (Fig. 176), S. eggersii (Fig. 177), s. flagellata (Fig. 178), S kraussiana (Fig. 179), S radrata (Fig. 180), S selaginoides (Fig. 181), S uncinata (Fig 182) * Fig 171. Selaginella firmula. Fragment of section through megaspore inembrane. x 2000.
J! LLLL Fig 172 Selaginella rupestris Fragment of section through megaspore membranc X 1000. Fig 173 Selaginella selaginoides. Fragment of section through megaspore membrane (* 2000). An electron micrograph showing the fine structure of part of the membrane is shown in PI III (facing P 94), lower figure. b Fig 174 Selaginella apus, microspore a, distal face; b), proximal fare (u, > x 1000). Lower left-hand cletail lateral view (Y 250); upper detail: selerine stratification (x 2000) 7-665017 a. Erdiman
A
?
Fig. 173. Selaginella alroviridis, microspore; sclerine stratification (* 2000)
and distal face (* 1000).
Intel
Fig. 176. Selaginella biformis, microspore. From left to right lateral view
(* 250), distal face (* 1000); cxine stratification (* 2500).
b
a
c
d
Fig. 177. Selaginella eggersii, microspore. A. distal (left) and proximal face
(right: surface and section); b), sclerine stratification; c, different types of
processes; d-f, aberrant microspores. - a, df x 1000; }),cx 2000.
95
b
Fig. 178. Selaginella flagellata, microspore; lateral view (* 250) and distal
(a) and proximal (b) face (* 1000)
b
t2
с
f
Fig 179 Selaginella kraussinna, microspore; a, proximal face (1000),
h, sclerine stratification (x 2000), c-e, successive patterns in LO-analysis
of the laesura, f, two microspores with coinmon cover (* 1000).
M
Fig 180 Selaginella radiala, microspore, sclerine slratification (2000)
and distal face (* 1000).
11 d b с Fig. 181 Selaginella selaginoides, microspore. Main figure. proximal face (* 1000), a, lateral view (* 250); is, sclerine stratification (2000); C, LOpatterns; d. part of lacsura at different adjustments from high (lent) to low (right). M Fig. 182. Selaginella uncinata, microspores in polar (leſt, distal face) and lateral view (right; Hoth * 1000). In the centre sclerine stratification ( x 2000). SELENODESMIUM: see Fig. 188 A, r. 96. STENOCHLAENA: Fig 183. Stenochlaena tenuifolia. From left to right: prorimal face (x 230); lateral view (surface and section; x 1000); sclerine stratification (* 2000).
PI III Lycopodium clavalum and Selaginella selaginoides Cpper figure; section through the evinc in Lycopodem cimatum An outer laminated and an inner granulated layer can be seen. IMG B V Afzelius (- 10,000). Lower figure section through part of the crine of a megaspore of Selaginella selaqınoides. ENG 13 V Afzelius (12,500) (From Siensk bot. Tidshr., vol. 48, 1954 ) 11. Fig. 181. Selaginella sclaginoides, microsporc. Main figure. proximal face (* 1000) a, lateral view (x 250); b, sclerine stratification (* 2000); <, LOpatterns; d, part of lacsura at different adjustments from high (left) to low (right). M Fis 182. Selaginella uncinata, microspores in polar (left, distal face) and lateral view (right; both 1000) In the centic. sclerinc stratification (2000). SELENODESMIUM: see Fig. 188 A, p. 96. STENOCHLAENA:- Fig 183. Stenochlaena tenuifolia. From left to right: proximal face (* 250). Jateral view (surface and section; x 1000); sclerine stratification (2000) 95 STICHERUS: see Fig. 114 B, p. 63 STROMATOPTERIS: see Fig. 114.1. p 03 TAENITIS:- g 184 Taemthis blechnoides From left to right sclerine stratification (2000) and lateral view (250); distal face (* 1000); LO-patterns TAPEINIDIUM:- ☺ Fug. 185. Tapeinidium pinnatum; proximal face (230), lateral view (surface and optical cross-section; y 1000); LO-patterns. THYRSOPTERIS: a, Fig. 186. Thyrsopteris elegans; proximal face (surface and optical crosssection; x 1000). Optical cross-sections through the laesura at ad are shown in agri,. e, LO-patterns. Lower left-hand detail. lateral view (* 250): upper right-hand detail: sclerine stratification (* 2000).
TMESIPTERIS:--
Fig 187. Tmesipleris forsteri, sclerine stratification (* 2000).
TODEA: see Fig. 155 F, p. 84.
TRICHOMANES (Fig. 188 A, B, D), HYMENOPHYLLUM (Fig. 188 C):
A
B
с
C
D
Fig 188. Hymenophyllaceae. A-C, Trichomanes and Hymenophyllum, cxinc
stratification (* 2000). A, T. (Selenodesmium) rigidum B, T (Afacroglena)
meifolium. C. Hymenophyllum (Hymenoglosslim) cruentum. D), Trichomanes
(Vandenboschia) radicans, proximal face (* 1000); a, cxine stratification
(* 2000); b, lateral view (* 250), C, LO-analysis of lacsura.
VANDENBOSCHIA: see Fig. 188 D.
VITTARIA:--
Fig. 189. Villaria vitlarıoides. From left to right: Jateral view (* 250); proxi.
mal face (* 1000); sclerine stratification (* 2000).
97
WOODSIA:-
Fig 190. Woodsia alpina. From left to right selerine stratification (* 2000),
distal face (* 1000), LO-patterns.
XIPHOPTERIS:
Fig. 191. Xiphopteris saffordit From left to right lateral sies (x 250), proti
mal face (surface and optical cross-section, * 1000); crine stratification
(* 2000).
opening several of these dry red Cases, I found them to be quite hollow, without any thing at all in them; whereas when I cut theni asunder with a sharp Pen-kniſe when green, I found in the middle of this great Case, another smaller round Case, between which two, the interstices were fill'd with multitudes of stringie fibres, which seem'd to suspend the lesser Case in the middle of the other, which (as farr as I was able to discern) seem'd full of exceeding small white seeds ..." From a description of moss capsules in R. I looke's Vicrographia 1667 (pp 131, 132) BRYOPHYTA Pl. IV (facing p. 108), V (facing p. 109); Figs 192-253. HEPATICAE Aneuraceae: Fig. 240 (p. 118). Marchantiaceae: Fig. 209 (p. 106). Anthocerotaceae: Figs. 194 (p 100), Monocleaceae Fig. 226 ( p113). 208 (p. 106), 225 (p. 112). Oxymitraceae: Fig. 230 (p. 114) Blasiaceae: Fig. 197 (p. 101) Pelliaceae: Fig. 231 (p 115) Cleveaceae: Fig. 201 (p. 101) Plagiochilaceae Fig. 233 (p. 116) Codoniaceae. Fig. 213 (p 108). Radulaceae Fig. 238 (p 118). Conocephalaceae: Fig. 205 (p. 104). Ricciaceae Fig. 211 (p 119). Corsiniaceae: Fig 206 (p. 105) Riellaceae Fig 242 (p. 120) Cyathodiaceae: Fig. 207 (p 105). Scapaniaceae: Fig. 243 (p. 121). Exormothecaceac: Fig. 211 (p 108) Sphaerocarpaceae. Fig. 216 (p. 109). Frullaniaceae: Fig. 214 (p. 109). Symphyogynaceae Fig. 250 (p. 123). Grimaldiaceae. Fig. 232 (p. 115). Targioniaceae Fig. 251 (p. 123). MUSCI Andreaeaceae: Fig. 192 (p. 100) Lembophyllaceae: Fig 228 A (p. 113). Archidiaceae: Fig. 195 (p. 101) Leptostomaceae: Fig 222 (p. 111). Bryaceae: Fig. 199 A-C, E (p. 102). Leskeaceae Fig. 223 (p. 111). Buxbaumiaceae. Fig. 200 A,B (p. 102) Leucobryaceae Fig. 203C (p. 103). Calomniaceae. Fig. 201 (p 103). Meeseaceae Fig. 224 A-C (p 112). Calymperaceae: Fig. 202 (p. 103). Mniaceae. Fig. 199 D (p. 102). Catascopiaceae. Fig. 221 D (p 112). Neckeraceae. Fig 228 B (p. 113). Dicranaceae. Fig. 203 B (p. 103). Oedipodiaceae Fig. 229 (p 114). Diphysciaceae. Fig. 200 C (p. 102). Orthotrichaceae: Fig. 253 (p 124). Ditrichaceae: Fig. 203 A (p. 103). Pleurophascaceae: Fig. 234 (p. 116) Encalyptaceae: Fig. 210 (p. 107). Polytrichaceae. Fig. 235 (p. 116). Ephemeraceae: Fig. 227 (p. 113). Pottiaceae. Figs. 193 (p. 100), 236 Fissidentaceae. Fig. 212 (p. 107) (p. 117). Funariaceae: PI IV, V. Rhizogoniaceae: Fig. 237 (p. 117). Georgiaceae: Fig. 215 (p. 109). Schistostegaceae. Fig. 211 (p. 121). Gigaspermaceae: Fig. 217 (p. 110). Sphagnaccae: Fig. 246 (p. 121). Grimmiaceae: Fig. 239 (p. 118). Spiridentaceae: Fig. 217 (p. 122). Hedwigiaceae: Fig. 218 (p. 110). Splachnaceae: Fig. 218 (p. 122). Helicophyllaceae: Fig 219 (p. 110). Symphyodontaceae: Fig. 249 (p. 122). Hookeriaceae: Fig. 220 (p. 111). Timmiaceae: Fig. 252 (p. 121). Hylocomiaceae: Fig. 221 (p. 111).
ACROSCHISMA:-
d
.
Fig. 192. Acroschisma wilsonii; a, distal face (* 1000); b, tetrad (* 1000),
C, LO-patterns (ilistal face); d, etine stratification (1, in chlorinated spores);
x 2000.
ANOECTANGIUM: -
BRUST
Fig 193. Anoectangium aestivum; sporc in lateral view, surface and optical
section (distal pole upwards; * 1000), LO-patterns (distal face)
ANTHIOCEROS:-
Fig 194. Anthoceros tuberculatus; from left to right: sclerine stratification
(* 2000), proximal face ( 825); LO-pattern.
ARCHIDIUM:
Fig. 195. Archidium Alternifolium, part of spore surface (* 1000)
ATHALAMIA:
Fig. 196. Athalamia nana, sporc (* 1000; and sclerine stratification (x2000),
BLASIA:
Fig. 197. Blasia pusilla; part of spore, surface and optical section (* 1000).
BLINDIA; see Fig. 203 B, p. 103.
BRACHIOLEJEUNIA:--
AAB
Fig. 198. Brachiolejeunia sandwicensis; part of spore, surface and optical
section (* 1000); sclerine stratification (* 2000) and LO-patterns,
BRYUM (Fig. 199 A, B), CINCLIDIUM (Fig 199 E), MIELICHHOFERIA
(Fig. 199 E), POHLIA (Fig. 199 C):-
A
B
EAVY
OD
E
Fig. 199. Bryaccac (A, B, C, E), Mniaceae (D) – A, Brytım mammillaluni;
part of spore, surface and optical section (* 1000). B. B. caespilicium;
spore in lateral view (* 1000) C. Pohlia elongata; spore in lateral view
(* 1000) D, Cinclidium subrotundunr; part of spore, surface and optical
section (* 1000) E. Vielich hoferia elongata; cf. proximal face (* 1000)
BUXBAUMIA (Fig. 200 A, B), DIPHYSCIUM (Fig. 200 C).
A
B
C
-
Fig. 200. Burbaumiales. A, Birbaumia aphylla. B, B. viridis. C, Diphyscium
foliosum. (A-C X1000).
103
CALOMNIUM:-
Fig 201. Calomnium laetum, surface and optical section (¥ 1000), exine
stratification (2000) and LO-patterns
CALYMPERES:
Fig. 202. Calymperes afzelii, surface and optical section (1000), exine
stratification (¥ 2000) and LO-patterns.
CATASCOPIUM: see Fig. 224 D, p. 112.
CERATODON (Fig. 203 A), BLINDIA (Fig. 203 B), LEUCOBRYUM (Fig.
203 C):--
B
A
d!
pr?
C
x 2000
Fig. 203 Dicranalcs – A, Ceralodon purpureus, from left to right: spore in
lateral view (* 250); two spores (cf. proximal face; X 1000), LO-patterns.
B, Blindia acuta; from left to right. spore in lateral view (* 250); of distal
face (* 1000); cf. proximal face (* 1000) C, Leucobryum glaucum; exinc
stratification (* 2000).
CINCLIDIUM: see Fig. 199 D, p. 102.
CLEVEA:-
Fig. 204. Clevea robusta, distal face (* 1000).
CONOCEPHALUM:--
Fig. 205. A, Conocephınlum (Fegalella) conicuns; part of spore (* 1000);
optical section and surface; LO-patterns. B, C. supradecompositum, part of
spore (* 1000), surface and optical section (with LO-patterns).
105
CORSINIA:
X 2000
d
*1000
250
* 2000
x 100
Fig. 206, Corsinia marchantiordes a, lateral view, b, sclerine stratification
in the less conver face; e, LO-pattern, d, sclerine stratification in the more
convex face; e, polar view (less conrer face), f1, fe part of spore in polar
view (fi, optical section, fs, surface)
CYATHODIUM
* 2000
Fig. 207. Cyathodium africanum; sclerinc stratification and LO-palterns
DENDROCEROS:-
Fig 208. Dendroceros crispalus; part of sporc surface (* 1000).
DIPHYSCIUM: see Fig. 200 C,
102.
DUMORTIERA:---
A
Fig. 209. Dumortiera hirsuta; exine stratification (2000) and LO-patterns
B, D. velulina, spore in polar view (* 1000), exinc stratification (* 2000),
and LO-patterns.
107
ENCALYPTA:-
A
pr?
di?
B
Fig. 210. 4. Encalypla alpina. From left to right. spore in lateral view ( *230);
cf. proximal lace (* 1000): 10.patlerns in the distal (?) face, 1, E. ciliata:
(listal face (upper ligure) and proximal face (lower figure: th fogo).
the lower right-hand detail shows a spore in lateral view (* 250). C, É.
rhabdocarpa; distal face (upper figure) and proximal face. (lower figure;
both x 1000); the lower right-hand detail shows a spote in lateral view
(* 250).
EXORMOTIIECA:
1 x 1000
x 250
x 250
50
100 M
Fig. 211. Erormotheca fimbriala; a, proximal, b, distal face (* 230) The
LO-patterns to the extreme right exhibit-at x-y--a part of one of the muroid
ridges shown in b, and also the "insulous" condition in the bottom of the
lumina
FISSIDENS:-
Fig. 212. Fissidens adiantoides; surface and optical
section (* 1000), evine stratification (* 2000), and
LO-patterns.
FOSSOMBRONIA:-
al
B
119
2
Fig. 213, A, Fossombronia angulosa; spore (Y 1000), 2, part of "muri" at
light and, in l», at slightly lower adjustment of the microscope: c, sporc,
jateral view (* 230); the LO-patterns to the extreme left show the patterns
at the bottom of the lumina. B, F. dumortieri; d, exine stratification (with
"murus", * 2000); e, spore (proximal face ?), * 250
109
FRULLANIA:-
Fig. 214. Frullania involvens; surface and optical section (* 1000)
FUNARIA: see Pl. IV, V (facing pp. 130, 131).
GEORGIA:-
Fig. 215. Georgia pellucida; from left to right spore in lateral view (x 250).
spore in polar view (* 1000), LO-patterns.
GEOTHALLUS:
di 7
pr?
CD
A
B
pr 7
Fig. 216. Geolhallus tuberosus; A, laleral sies (* 230), B, polar vies (di
cf. distal face, surface and optical section: pr = cf. proximal face, surface,
* 250), C, selerine stratification (about x 2000).
GIGASPERMUM (Fig. 217 B), LORENTZIELLA (Fig. 217 A):- B A Fig. 217. Gigaspermiaceae. A, Lorentziella paraguensis; sclerinc stratification (* 2000) and LO-patterns. B, Gigaspermum repens; sclerine stratification (* 2000) and LO-patterns HEDWIGIA (Fig. 218 A), HEDWIGIDIUM (Fig 218 B).-- А. for Fig. 218. Hedwigiaceae - A, Hedwigia ciliata; from left to right: proximal face (* 250); spore in lateral view (* 1000); LO-patterns. B, Hedwigidium integrifolium, from left to right proximal facc (* 250), spore in lateral view (* 1000), LO-patterns. (From Erdtman in Svensk hot Tidshr 1954.) HEDWIGIDIUM: see Fig. 218 B. HELICOPHYLLUM:- ab Fig. 219. Helicophyllum forquatum; a, LO-patterns, proximal face; b, LOpatterns, distal face; the central figure shows the proximal face of a sporc, surface (left) and optical section (right; X 1000).
111 M HOOKERIA:- 1 Fig 220. Hooleria albicans, spore (X 1000) and LO-patterns. IIYLOCOMIUM:- Fig. 221. Hylocomium splendens, the main figure shows the supposed proximal spore face (1000) ISOTHECIUM: see Fig. 228 A, p. 113. LEPTOSTOMUM: Fig. 222. Leptostomum macrocarpum, from left to right. spore in cf trans- Verse lateral view (* 250), spore in longitudinal lateral view (x 1000); LOpatterns. Upper detail figure erine stratification (* 2000) LESKEA: Fig. 223. Leshea polycarpa; spore, surface and optical section (* 1000) The left part of the right-hand detail Gigure shows the LO-patterns of the cf. clistal face, the right part those of thic cl. proximal face.
LEUCOBRYUM: see Fig. 203 C, p. 103.
LORENTZIELLA: see Fig. 217 A, p.
110.
MEESEA (Fig. 224 A, B), CATASCOPIUM (Fig. 224 D), PALUDELLA (Fig.
224 C):
A
C
B
D
pr
dr
Fig. 224. Meeseaceae (A-C), Catascopiaceae (D). - A, Veesea longisela, cf.
distal face (left) and cf. proximal face (right; 1000), B, M uliginosa, cf.
distal face (left) and cf proximal face (right; * 1000). C, Paludella squarrosa,
cf. distal face (len) and cf. proximal face (right; ~ 1000) D, Catascopinn
nigritum, part of sporc (* 1000), surface (left) and optical section (right).
MEGACEROS:
b
Fig. 225. Vegaceros aosanus; a, spore in lateral, slightly oblique view (x1000);
b, distal face (* 250); c, elater (* 1000, the detail figure shows the wall
stratification, X 2000).
MIELICHHOFERIA: see Fig. 199 E, p. 102.
113
MONOCLEA:
L
Fig. 226. Monoclea forsteri; spore (1000), euine stratification (* 2000),
and LO-patterns.
NANOMITRIUM:
x 500
b
Fig. 227. Nanomilrium tenerum; a, laleral view (x 250), b, outline of tetrad
(* 250); c, proximal face, surface and optical section (> 1000); d, distal
face (< 500).
NECKERA (Fig. 228 B), ISOTHECIUM (Fig. 228 A):-
SAN
d
B
Fig. 228, Neckerinales. - A, Isothecium myosuroides; LO-patterns and spore
(* 1000). a, surface; b, optical section (upper quadrant without, lower
quadrant with, easily detachable "granules'). B, Vechera complanata; sporc
(* 1000), surface and optical section; exine stratification (* 2000), and
LO-patterns.
OEDIPODIUM:
0
Fig. 229. Oedipodium griffithianum, a, proximal face, surfacc and optic
section (* 1000), b, distal face (* 250); C, lateral vicw (* 250); d, LC
patterns, e, sclerine stratification (about x 2000).
C
OXYMITRA:
b
Fig. 230. Orymitra paleacea; a, proximal (?) face, surfacc (left) and section
(right; * 1000); b, spore in lateral view (250); c, optical section through
à murus" in the reticulum, d-6. LO-analysis of a 'muriferous" part of
the spore surface.
9
PALUDELLA: see Fig. 221 C, p. 112.
115
PELLIA:
Fig. 231. Pellia epiphylla; part of a spore, optical section and surface ( *1000);
erine stratification (* 2000)
PHASCUM: see Fig. 236 A, p. 117.
PLAGIOCHASMA:-
b
Fig. 232. Plagiochasma intermedium; a, proximal face (* 1000) and LO.
patterns; h, one half of an elater (* 1000)
116
PLAGIOCHILA:
AL?
Fig 233 Plugtochila asplentodes; spore (surface and optical section, 1000);
LO-patterns.
PLEUROPIASCUM:-
Ő
b
@o
@
Fig 234 Pleurophascum quadrilobum; main figure proximal facc (surfacc
and optıcal section; * 1000); a, proximal face (* 250) of slightly irregular
spores; b, transverse side view (* 250); c, longitudinal side view (* 250,
distal pole upwards).
POHLIA: see Fig. 199 C, p 102.
POLYTRICIUM:-
B
b
Fig. 235. A, Polytrichum juniperinum (* 1000), a, spore in lateral view
(proximal pole upwards); b, prorimal face; c, distal face and LO-patterns
of the square shown in the centre of the face. B, P. gracile; d, lateral view;
e, proximal (tenuitatiferous) face; f, distal face; &, fetrar (* 250; d-f * 1000),
117
POTTIA (Fig. 233 B), PRASCUM (Fig. 233 A);-
A
B
1800
E
Fig. 236. Pottiaceae (see also Fig 193, P 100) A, Phascum cuspidalun,
from left to right spore in polar view (* 250); spore in lateral view, surface
(left) and optical section (right, x 1000), LO-patterns B-E, surfacc (*1000,
only a fourth of a hemisphere shown), sclerine stratification (about x 2000),
and LO-patterns in Pothin crinita (B), P davallana (C), P. Iruncata (1)), and
P. heimii (E).
PYRRHOBRYUM:--
Bts
Fig 237. Pyrrhobryum spiniforme. From left to right: spore in lateral siew
(* 250); spore in polar view (* 1000), surface (left) and optical section;
LO-patterns.
RADULA:
Fig 238. Radulo lindbergiana; surface and optical section (lower right-hand
quadrant, * 1000); crine stratification (* 2000), and LO-patterns.
RHACOMITRIUM:--
Fig 239 Rhacomtrium microcarpon From left to right: spore in lateral view
(* 250), spore in polar view (* 1000); LO-patterns
RICCARDIA---
Fig. 240. Riccardia latifrons, spore (* 1000); erine stratification (* 2000);
LO-patterns.
119
RICCIA:
d
ad
Fig. 241. Riccia beyrichiana, main figure; distal face, surface and optical
Şection (left); proximal face, surface (right, ~ 1000); a, laesura (?); b. spore
in lateral view (x 230); c, outline of sclerine stratification (without attempt
to classify the layers); d, sporc margin (optical section); x-y, approximate
Jimit between listal and proximal face.
RIELLA: Fig. 242. A, Riella halophila (left). B, R. purpurcospora (right). The sectors in the main figure (A + B) are as follows enumerated clockwise, beginning at A): sclerine, optical section: part of proximal (?), tenuitatiferous (7) face: part of distal (?) face; sclerine, optical section, part of proximal (?), tenuitatifcrous (?) face; part of distal (Y) face. In the lower left-hand detail figure tentative interpretations of the selerine stratification in R. halophula arc el hibited. The lower riglit-liani delail shows an outline of the sclerine stratification in R. purpureospora (the second layer from below should have been blackcned in, not dotted), the Ol-pattern is due to reticuloid arrangement of the 'sexinous' (cf. endosexinous) ciements in the third layer from below.
121
SCAPANIA:
Fig. 243 Scapania paludicola, part of spore surface
(* 1000), LO-patterns
SCHISTOSTEGA:-
00000
Fig. 244 Schistostega osmundarea,
LO-patterns and five spores (Y
1000, from left to right optical
section, distal face of three spores.
and spore in lateral view), the
upper detail figure exhibiting the
exine stratification is enlarged
about 4000 times
SOUTHBYA
b
Fig 245 Southbya stillicidiorum; a, cf, distal
face, b, cf proximal face (* 1000)
SPHAGNUM:--
A
B
C
Fig. 246. A, Sphagnum fimbriatum, les vating (tetrachotomolaesurate" or
*4-letc") spore (* 1000) 1. S. palustre: proximal face of normal (3-lete)
spore (* 1000), a, with perine (surface), b), with perine (optical section);
c, without perine (surface)
12 SPIRIDENS: C17) D Fig. 247. Spiridens aristidifolius; sporc in lateral view (* 1000); crine stratification (2000). SPLACHNUM (Fig. 248 C), TAYLORIA (Fig. 248 B), VOITIA (Fig. 248 A). B А. Fig. 248 Splachnaceae. -- A, Voilia nivalis (* 1000) B, Tayloria lingulala: surface and optical section (* 1000), LO-patterns, and exine stratification (* 2000) C, Splachnum vasculosun, two bilateral spores (the upper in lateral view, proximal face upwards; the lower in polar vicw, showing the cf. proximal tenuitas) and one # radiosymmetric sporc (* 1000). STROEMIA: see Fı: 253 A, p. 121. SYMPHYODON:- Fig 249. Symphyodon echinatus; spore in lateral (?) view (* 250); spore in polar (?) view (* 1000); LO-patterns.
123 SYMPHYOGYNAS Fig 250. Symphyogyna podophylla, four spores (three 1.0-patterns 250, one < 1000); TARGIONIA:- Fig. 231. Targionia hypophylla, from left to right: spore in lateral view (* 250); spore in polar view (1000), selerine stratification and! LOpalterns. TAYLORIA: see Fig. 248 B, p. 122. 9- KR51
TIMMIA:-
B 229
А.
Fig. 252 A, Timmia austriaca; spore in lateral view (* 1000); LO-patterns.
B, T anomala, exine stratification (* 2000)
ULOTA (Fig. 253 B), STROEMIA (Fig. 253 A);
A
B
a
Fig. 253. A, Stroemin gymnosloma, a, spore in lateral view (* 250); b, distal
face (* 1000), c, proximal face (* 1000), d, tetrad (* 250); c, outline of
apertures. B, Ulola bruchit; surface and optical section (* 1000).
VOITIA: see Fig. 248 A, p. 122.
SUPPLEMENT ON NEW METHODS IN PHYSICAL CELL RESEARCH AND THEIR APPLICATION IN STUDIES OF POLLEN GRAINS AND SPORES BY BM. AFZELIUS The rapid development in physical cell research has recently provided much news of interest to pollen and spore morphology. Deeper understanding of the cell, the morphology and composition of its organellae and their finest constituents down to macromolecular dimensions can be gained through the ever-widening application of refined techniques such as phase contrast and polarization microscopy, ultra-violet light, interference, and electron microscopy. Certain indirect methods are also of interest (e.g. X-ray diffraction) and there are microscopical and other methods for obtaining quantitative cytochemical data. Micrographs of macromolecules can thus be made (cf. Wyckoff 1919, p. 189) and it is even possible (cf. e.g. Frey-Wyssling 1954) to determine the macromolecular or particle weight by microscopical methods. From a study of electron micrographs (Steinmann in Finean, Sjostrand, and Steinmann 1953) of the small spherical particles constituting the lamellae of chloroplast grana Frey-Wyssling was able to calculate their size as being equal to one to four Svedberg units, containing eight to twelve chlorophyll molecules (the molecular weight of one Swedberg unit - 17,600) Moreover, micrographs showing the exact localızation of enzymes in the cell are now available (Sheldon, Zetterqvist, and Brandes 1955). By means of microscopical interferometric methods it is, furthermore, possible to determine mass as small as 001 ps/11* (Davies, Engström, and Lindstrom 1953; 1 pg -- 10-12 g). Microspectroscopical methods allow of determination of mass as small as 0.001 pg (Caspersson 1950). To be able to make use of the new developments in the field of microscopy it is often necessary to introduce new methods for preparing the materiat to be studied. Thus the invention of the electron microscope (Knoll and Ruska 1932) called for intense research for new methods of preparation. As far as biology is concerned satisfactory results were not achieved until ultra-thin sections could be made by routine methods-almost 20 years after the invention of the clectron microscope. Several new ways of preparation might be convenient in ordinary light microscopy but ultra-thin sectioning will, no doubt, eventually be the most useful one. În response to a request of visitors to the Palynological Laboratory, Stockholm-Bromma, some of the problems met with when using the ultrathin section technique wall be discussed in the following. Alethods of preparation For studies of ultra-thin sections of non-acetolyzed pollen grains and spores it is, i.a., advisable to consider the question of fixation carefully. For light microscopical work fixation by means of formalin, or freezing and drying is often sufficient. Fixation with osmic acid is, however, to be preferred; in fact, in electron microscopical studies it is, so far, the only reliable fixative. Osmic acid also acts as an electron stain. In acetolyzed exines the staining effect of thc osmic acid is negligible and probably of no usc. Experiments ought to be made to discover, if possible, other heavy metals or ions suitable as electron stains. In investigations of mammalian tissues a buffered solution of osmium tetroxide is usually used (acetateveronal buffer at pH 7.2; cf. Rhodin 195-1). The same fixative has been used by Turian (1956) and Afzelius (1955). In the opinion of P. Sitte (personal communication, November 1956), buffering of the solution is not particularly important when fixing plant cells which have not formed a large vacuole, since the buffer might penetrate the cell morc rapidly than the osmic acid and thus cause a poisoning of the cell before fixation. Sitte, moreover, emphasized the importance of using an isotonic solution of 1 per cent osmium tetroxide by adding sugar or urethan. The slow penetration of osmic acid presents a problem familiar to all who have tried this type of fixation. In pollen grains the fixative probably penetrates into the grains more readily through the aperture membranes than through the wall proper. (N.B. In order to obtain good fixations to meet the high demands of electron microscopy it might be advisable to cut large pollen grains or spores before fixing). Wettstein has obtained very good results by using osmuum vapour (cf. Pl. IV, V). The necessity of using suitable embedding media in thic study of thin sections hy mcans of phase contrast has recently becn stressed. Ordinary histological techniques have, as a rule, been adjusted to thic use of much thicker sections, for which glycerol jelly is often a good embedding medium. Thin sections (about 500-5000 A thick) should preferably he studied in water, ultra-thin sections (thickness < 500 A) in air (water and air are better than glycerol jelly on account of their lower refractive index). The following formula shows the importance of a wide difference between the refractive index of the object and that of the embedding medium when the geometrical thickness of the section is decreased OPD=ean, -ng); OPD, optical path difference (see p. 131), Ng, refractive index of the object; n,, that of the enclosing medium, e, geometrical thickness of the object. If air is used as an enclosing medium it is, of course, possible to apply cover slips and immersion oil in the ordinary way For theoretical reasons, however, full use cannot be made of the numerical aperture of the microscope Good results have, nevertheless, been obtained by this method. When specifying the thickness of the sections, e.g ultra-thin sections embedded in air, the figures originally obtained from electron micrographs of shadowed sections (Sjostrand, personal communication) have been used. The present author has measured the thickness of ultra-thin sections by means of interference microscopy. These measurements, in general, tend to give slightly higher values than those calculated from shadowed sections. It may perhaps not be superfluous to point to another circumstance that might cause confusion. If thin exine sections are studied by means of an ordinary light microscope a number of diffraction lines can usually be seen, due to superposed light waves in the thin exine The number of the waves can be altered by changing the wave length of the light used (Ingelstam, personal communication). The diffraction lines must not be mistaken for lamellae or demarcation lines between different layers of the exine as has apparently been done by Pflug in describing optical sections of entire grains (cf. Pflug 1953, p. 66. "Exospore and Exinen sind aus konzentrischen Schalen (Lamellae) zusammengesetzt, das ist in unzahligen Fallen beobachtet. Fraglich ist nur deren Zahl. Sie ist einmal subjektiv bedingt durch die Scharfe der Optık"). Ultra-thin sections should always be properly stained in order to obtain as much information on wall stratification, cell contents etc. as possible. The fact that the immature wall of certain fungus spores is easily penetrated by stains (Nannfeldt, personal communication) seems to open up a possibility for obtaining further details on sporoderm formation. Further research work in sporodermi morphogenesis is indeed necessary. In this context another problem may be mentioned: electron micrographs of acetolyzed exines often show that the exine, the endonexine excepted, is altogether homogeneous-amorphous (cf. e.g. EMG by Afzelius in Érdtman, 1956 a, Fig. 1, p. 132) No demarcation lines can be observed at a resolution of 40-50 A. Oshurkova (1936) in a review of a paper on the fine 128 structure of the pollen wall in Clivia miniata (Afzelius 1955) criticizes the use of several terms to indicate different parts of this continuous layer. I attention is paid to fine structure only this criticism seems to be justified in the case of Clivia, and many other pollen types. There are, however, exceptions. The basal part of sexinous rods in pollen grains of Cobaea penduliflora (Erdtman 1952), Cucurbita (Muhlethaler 1955), Linum, and some other genera (Erdtman 1956 b) is fixed well into the nexine. Studies of ultra-thin sections, suitably stained, would probably shed more light on these and similar problems, and make possible further contributions to the elucidation of the morphogenesis of the sporodcrms. Electron microscopy There are practically no pollen grains or spores small cnough to make possible a study of unsectioned grains or spores in the clectron microscope. However, some efforts along these lines have in fact been made (cf. eg. Burton 1916). Ueno (1919, and personal communication 1956), isolated "viscin threads" of Oenothera odorala and Gaura lindheimeri etc., and studied them, without further preparation, by means of clectron microscopy'. These threads, or part of them, probably consist of the same material as the exine, since they are not dissolved if heated in an acetolysis mixture (cf. Erdtman 1952, Fig. 170 B:b, p. 292, and Fig. 171A, p. 293). Fragmentation of material, by means of ultra-sonics or other methods, was often carried out before the introduction of ultra-thin sectioning techniques (cf. the spine of a malvaceous pollen grain, fig. 151A, p. 262, in Erdtman 1952). It is a rapid method of preparation, but rather rough and unsatisfactory. Many fragments will be too thick and thus instantly charted by the electron beam; they defile the microscope and the method cannot be used in high resolution microscopy. Although now outmoded in clectron microscopy, this technique can still be of some use in ordinary microscopical investigations. The interpretation of the micrographs is easier if the material consists of one or a few structural units only, as when dealing with acetolyzed exines. Replica methods are often useful in the study of the surface details of pollen grains and spores. Sitte (1933) has modified the wax impression method by Mahl (1912), whereas Bradley (1954, 1937), and Mühlethaler (1955) have used, with much success, a carbon replica method derised by Bradley. Bradley has thus provided new information concerning the colpi in the pollen grains of Lamium album; he has also found that the spores Russula verrucosa are covered with a thin amyloid film. of 120 Fragmentation and replica-making can be combined with shadowing by means of metal-evaporation, although this treatment is ton rough in studies of cytoplasm ctc. (silver, or aluminium, is often used in light microscopy, palladium in electron microscopy) The thickness of the metal film must be calculated with care, as must also the angle at which the metal atoms are to strike the material (cf. micrographs in Wyckoff 1919, p 110, 121, and 227–232). In high resolution electron microscopy the method has now almost lost its importance since sufficient contrast is preferably obtained by using suitable electron stains, A particularly penetrating evidence concerning the structure of the wall will, no doubt, ultimately be derived from studies of ultra-thin sections. Various types of fine structure in the resistant part of pollen and spore walls have recently been described by various authors. At a resolution of about 30 Å (cf. EMG, Figure 142, page 77) acetolyzed sporoderm seems to consist of small granules, 50-60 Å in diameter They are either arranged in lamellae, with the granules in a single layer, or grouped into an amorplious type of structure. The lamellate fine structure can be transformed into the latter type of structure by means of oxydation, as has been proved in Lycopodium clavatum (Afzelius 1956). Hitherto, attention has mainly been centred on the fine structure of the exine Electron microscopy may, however, also help to reveal the fine details in non-acetolyzed sporoderms and the interior of pollen grains and spores, as shown by some electron micrographs by D. von Wettstein reproduced in Pl. IV and V The following constitutes von Wettstein's description and interpretation of his EMG.S The spore wall of Funaria hygrometrica is divided into three layers, intine, inner exinc, and outer exine, each of which is structurally homogeneous and cannot be subdivided. The intine consists of a three-dimensional framework of fibrillae (probably cellulose). In properly fixed material the cytoplasm of the spore is in close contact with the intine and obviously not separated from it by a membrane (Pl. IV, Fig. c). The outermost layer of the cytoplasın seems to consist of a framework of fibrillae related in size, but not in chemical composition, to that of the intine. If, in the course of fixation, the cytoplasm is separated from the intine, some sort of membrane appears in the micrographs. The inner layer of the exine in Funaria displays a higher electron density than the intine. A kind of fibrillar structure seems to be present although obscured by a substance between the fibrillac. The intine/exine limit is marked by a fine and dense zone about 50-100 A in thickness (Pl. IV, Fig c). In Pl. IV, Fig. b, the exinc secms to be separated from the intine by a special, comparatively thick, layer. Olviously, a corresponding layer is lacking in Figs. a and c in the same plate. These, however, exhibit radial, strictly perpendicular sections through the wall, whereas Fig. b exhibits an oblique section, as can be inferred from the greater thickness of the exine as well as of the intine. Thus the effect of a special layer between exine and intine is produced by the high focal depth of the clectron microscope. In reality, this layer is nothing else than an overlapping of the intine and exine, due to oblique cutting. The outer part of the exine consists of a thin layer provided with large processes. It shows a high electron density and no inner structure could be resolved so far. The processes are finely papillate and exhibit a considerable variation as to size and shape. Tangential sections through the outer part of the exine (Figs. d-e) reveal the presence of a thin superficial layer (about 150-300 A thick) covering the processes as well as the rest of the exine surface. This layer has an extremely high electron density. The inner (infrasporodermal) part of the spore consists of cytoplasm, organellac, and reserve substances. Two kinds of organellac, viz. the mitochondria and the plastids, are easy to identify: the former contain a folded structure, which is also encountered in other plants and in animals, the plastids are considerably larger and provided with some traces of lamellae and globular particles. They lack the typical structure found in fully grown and assimilating chloroplasts. In Pl. IV, Fig. a, three chloroplasts and part of a fourth can be seen. The membranes surrounding the plastids display an interesting structure. In the last-mentioned figure there are many other small bodies with a slightly irregular surface, possibly reserve substances. They appear to consist of two components, a central grain with small clectron scattering power (white in the picture), and a surrounding layer of electronoptically dense and homogeneous material. There is reason to believe that these particles constitute a polysaccharide enclosed in a fatty substance. As can be concluded from this description, a tangential section gives a more or less uneven "black" area in the pictures. Organellae and inclusions of other kinds are depicted in Figs. f-i, amongst which are a great number of droplets with a strong electron scattering power. The other three organellae in these figures differ in size and structure. They have a special type of surface membrane which is composed of a thin layer supported by densc droplets, giving, in cross-section, the impression of a pearl necklace. The chemical constitution and function of these organellae are unknown. In Paris, in 1954, Frey-Wyssling, when summing up the facts known about the submicroscopic structure of the plant cell, intimated that the fine structure of the ground plasm was influenced by the fixative used but was still largely unknown. He further intimated that the presence of inicrosomes and mitochondria had been proved, but that a lamellate structure similar to that demonstrated in the large mitochondria of the hidney had not yet been found. Neither was it possible to ascertain iſ the nucleus was provided with a true nuclear membrane or not. The last two questions non *, GCM с !! IV l'unaria hygrometrus CMG D von Wellstein a, section through part of a spore, \, mitochondria, P, plastid 23,000 - ), oblique section of Sporoderm (intine and two-lavered exine) 334,000 --C, radial (perpendice ular) section through poroderm, csloplasmn, intine, and esine closely connected 31,000
Pl. V
000
09
h.
P. V. Funaria hygrometrica CMG D. von Wettstein , outer part of evine
(tangential section)
* 12,000
-C, same as dd; note the fine structure of
the interior, main part of the processes * 23,110/6). - l-i, parlich in the
cytoplasm {, h, 233,000, : * 17,000
131 seem to be answered in the affirmative by Turian kellenberger (1976). whose recent studies, together with those of von Wertstein, are certains the most penetrating analysis of plant (ell ultra-si cure hnown to the present author. Polarization microscopy, X-ray diffraction, Ul-microscopy and their possible application Polarization microscopy scems to be of little use in the study of acetolyzed erines; a form birefringence has not been revcaled and it has equally not been possible to trace the fine lamination of the exine in Lycopodium in this way. Probably the lamellate layer is too thin to provide an cffect in the polarization microscope. Freytag (manuscript Sept. 1956) has shown that an outer non-sporopolleninous layer can be demonstrated in some pollen grains by means of polarized light (cf. also electron microscopical studies of a similar layer by Afzelius (1955) and Bradley (1956)]. Likewise, X-ray diffraction does not provide any information about the exinc and, therefore, seems to be of no use for obtaining information on the fine lamination in Lycopodium clavatum. This is possibly due to the fact that the laminated layers are not parallel but curved and undulating (Engstrum, personal communication 1955). Engstrom has investigated dry, nonacetolyzed pollen grains of Alnus glutinosa and spores of Lycopodium clavalum by means of low angle diffraction without obtaining positive results. On the other hand, Labouriau and Cardoso, when investigating Lycopodium Prines by means of wide angle diffraction patterns, encountered a crystalline structure, a fact wluch seems difficult to explain (Acad. Bras. Ciênc. 1918). UV-optics have chiefly been used in microspectroscopical investigations. Wilkins (1953) describes the absorption difference between the two types of nuclei in the pollen grains in a UV-micrograph obtained by mcans of reflection microscopy, Interference microscopy The light passing through small objects in a microscope will be delayed in comparison to that passing through the surrounding merum, if the object, as is generally the case, has higher refractive index than the sarrounding medium. The interference microscope aims at measurements of the "optical path difference" (OPI), i.e., the difference in optical path through the object and an equal thickness of the surrounding medium, 104 e.g, water). When the OPD is known, the dry mass (m), as shown by Davies, Wilkins, Chayen and La Cour (1951), can be calculated from the following formula: 0.A m= 2 lo • OPD in water; A =area cms, % for a dry object can be calculated from the formula No1 where ea is the density of the object, no is the refractive index of the object, and no that of water. In cell substances y usually varies between 0.14 and 0.19. A x value of 0.18 will give dry weight determinations with an error not exceeding 10%. If the cell consists of a substance of one kind only, the error will be greater; in a mixture it will be less. In cells with plenty of storage products, e.g. many kinds of pollen grains and spores, it may not be advisable to use the average value of x. The thickness of the preparation or cell (1), is calculated from the formula P. = (no-nult All interferometric methods provide about the same accuracy. The OPD measurements of homogeneous, phase-even objects can be made with a precision of +20 A (+2 x 10-7 cm). In measurements of heterogeneous objects the same precision cannot be attained. Due to diffraction effects measurements of areas smaller than about 5 42 are not advisable. It is thus possible to make determinations of dry weight of 0.05 pg (0.01 pg/). In order to get the best resolution the Dyson interference microscope and that by Baker have a condenser with a large numerical aperture. The light traverses the object obliquely and the measurement values therefore tend to be slightly too high (Ingelstam and Johansson 1957). In another interference microscope described by Johansson and Afzelius (1956) and Johansson (1957), parallel light, perpendicularly passing the object, is used. Thereby a greater precision in OPD measurements is achieved. Measurements of the mass difference at various stages of pollen grain development, as well as at various stages of enzymatic digestion of pollen grains have been made by Davies et al. (1951). Measurements of the OPD of the exine, intine, cytoplasm, and nucleus of pollen grains have also been made (Johansson and Afzelius 1956). Besides the interferometric methods, quantitative cytochemical data may be obtained in other ways. Engstrom (1951) and Engstrum and Lindstrom (1917, 1950) have measured the mass of cells by means of soft X-rays. The method has also been tried on pollen material by Dahl and Engström (1951). Davies, Engström, and Lindström (1953) comment upon the X-ray absorption method and make comparisons with interferometric ones Using the X-ray technique measurements of 0 1 pg/12 can be made. The Dyson interference microscope provides measurements one order of magnitude greater (Davies et al. 1953), that is 0 01 pg/m2 Furthermore, it is possible to apply the interference microscope in investigations of living material. G. Bahr (1957) has succeeded in using the electron microscope for quantitative cytochemical measurements. As an example it may be mentioned that his calculation of the weight of a single microsome gives an approximate value of 8 to 16 x 10-19 g. General methodological problems There is no longer a tendency to separate form and function but to try to coordinate understanding of both in order to obtain an integrated picture of the living cell. It is, however, not an easy task to evaluate data obtained by different direct and indirect methods and to combine them to form a conception of the cell. It must also be borne in mind that different results are occasionally obtained even when applying the same technique to identical material. Likewise certain EMG.s may be interpreted in different ways by different workers etc. To my mind no purpose is served in trying to reconcile results and interpretations which do not fit; at the present state of knowledge and technique it is better to accept and acknowledge the divergences. But such problems, especially those arising from differences in interpretations, lead beyond the more particular problems of our field into those of a more methodological/epistemological character. 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Nature, Lond 172, p 1011 DAVIES, II G, WILKINS, M H , CITAYEN, J, and La Cour, L 1951 The use of the Interference microscope to determine dry mass in living cells and as a quantitative cytochemical method Quart J Micr. Sci 95. 3 101 Dysov, J 1950 An interferometer microscope. Proc. Roy Soc. Lond. A, 204, p. 1077. ENGSTRON, A 1946 Quantitative micro- and histocheinical elementary analysis by roentgear absorption spectrography. Acla rudtol., Suppl 63 ENGSTROV, A and LINDSTROY, B 1917 Histochemical analysis by X-rays of long wavelengths Erperientia 3 5 1950. A method for the determination of the mass ol extremely small blological objects Diochim biophys Acta 4-6 CROTVAN, G 1952 Pollen morpholngy and plant taxonomy | Angiosperms. Uppsala. 1956 (a) Current trends in palynological rescarch work Grana palynologica 1 2 1956 (b) "LO-analysis" and "Welcker's rule", a centenary Svensk bel. Tulshr. 50.1. 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Stockholm (Thesis ) SHELDON, HI , ZETTERQVIST, 11 and BRANDES, D 1955 Pistocherical reactions for electron microscopy, acid phosphates Exp Cell Res 9 3 SITTE, P. 1953 Untersuchungen zur submikroskopischen Morphologie der Pollest- und Sporenmembranen Akroskopie 8 9/10 SSOSTPAND, F. S. 1933 A new microtome for ultra-thin sectioning for high resolution microscopy. Erperientia 9 3 TURJAN, G and KFLLC\BERGER, E 1936 Ultrastructure du corps paranucléaire, des mitochone dries et de la membrane nucléaire des gamètes d'Allomyces macrogynus Erp Cell lies 11. UENO, J. 1919 (On the structure of visein threads as revealed by the electron mileroscope) Kagala 19 7 (In Japanese ) WETTSYFIX, D yov, 1957 Chlorophyll-Letale und der submikroskopische Formwechsst der Plastiden. Exp. Cell Res 12 3 II (LKINS, MH F 1933 The performance of spherical-mirror reflecting objectives when used for ultra-slolet photomicrography. J Noy. Microscop Soc. 73 2 Wyckoff, B. W. 1919. Clectron microscopy. Teclinique and applications New York and London.
ON THE CUTTING OF ULTRA-THIN SECTIONS BY J. RADWAN PRACLOWSKI Preliminary treatment of pollen or spore material and filling of gelatine capsules with methacrylate Before embedding in methacrylate, the pollen or spore material to be sectioned is successively washed in a centrifuge tube with distilled water (three times), 95 per cent alcohol (twice), and absolute alcohol (three times) The material should remain in the last-mentioned fluid for 21 hours. The alcohol is then decanted and some methacrylate (methyl methacrylate one part, butyl methacrylate seven parts) is poured into the tube. (X. B Liquid methacrylate contains a stabilizer which impedes polymerization and must be washed away with a two per cent NaOH solution before the methacrylate is poured into the tube ) The methacrylate is renewed three times at intervals of two hours and should be shaken, often and thoroughly, in order to prevent the pollen grains from sedimenting. After the last re-filling the fluid is shaken once again and transferred by means of a small suction pipe to gelatine capsules (No. 3 from Parke, Davis & Co., London, sec Fig. 251). It is important to add a proper amount of pollen grains ot the methacrylate. If too many grains are added the polymerization of the methacrylate (vide postea) may be rendered more difficult. Polymerization in UV light The liquid methacrylate is polymerized, i.e., transferred into a solid state by exposition to UV radiation; this method is to be preferred to polymerization induced by heat with the aid of a catalyzer, since the catalyzer may oxidize the sporopollepine. The filled gelatine capsules are covered with lids and fastened in pairs to a length of tape. About ten capsules to every tape are fastened in this way and exposed to the light produced by a 120 watt UV lamp. The distance between the lamp and the tape should be about SUCTION PIPE GLASS TUBE LON- TAINING POLLEN IN LIQUID METHACRYLATE. 2 GELATINE CAPSULES FILLED WITH METHA- CRYLATE. GELATINE CAPSULE WITH COVERING LIO (ORDINARY SIZE) CARDBOARD HOLD- ER IN WHICH THE GELATINE CAPSU- LES ARE PLACED READY FOR FILLING THU Fig. 251. Outfit for filling the gelatine capsules 15 cm. At the Palynological Laboratory, Bromma, the gelatine capsules are usually exposed to UV radiation during five nights, approximating 15 hours per night. During the first night, before the hardening of the plastic, the heat from the UV lamp should be checked by a ventilator in order to avoid the formation of air-bubbles in the sediment at the bottom of the capsules before the methacrylate solidifies (Fig. 255). After about 80 hours of exposition to the UV lamp the plastic is, as a rule, suſſiciently hard for cutting, Cutting is rendered difficult if there are air-bubbles in the methacrylale. The bubbles may be due to various causes, for instance, as already men- 137 HANGING LENGTH OF TAPE WITH GELATINE CAPSULES VENTILATOR UKLAMA Fig. 255 Polymerization of the methacrylate with UV light. tioned, to over-heating during the initial stages of polymerization, or to an over-concentration of pollen grains, or insufficient washings of the pollen material with methacrylate A capsule with air-bubbles can, nevertheless, be used for cutting of the pyramid (sec below) is made either at the side of the bubble or just above it. The cutting of the pyramid After soaking in water the gelatine capsulcs can be casily removed from the plastic units which, when dry, are ready for cutting. A unit is fixed in a steel object-holder (made by LKB-Produkter, Stocholm 12) in such a way that three to four millimetres of the pollen-learing part protrude from its mouth (Fig. 236). After trimming the top of the plastic unit with a razor blade (Fig. 257), a thin foursided process, the pyramid, containing the pollen grains or spores for sectioning is carved from the apex.
PLASTIC UNIT 30mm. -> SIDE VIEW OF LKB HOLDER MADE UP OF 2 INDEPENDENT PIECES OF STEEL MOUNTED AND SCREWED TOGETHER (WITH A KEY). De VIEW FROM ABOVE OF THE LKB HOLDER WITH MOUN- TED PLASTIC UNIT. mm. Fig 256. LKB object-holder with plastic unit mounted (1:1). When estimating the size of the pyramid, the following three factor should be taken into account: 1. the desired thickness of the sections (the thicker the sections the larger, and especially the higher, the pyramid must be), 2. the size of the pollen grains (the size of the pyramid should be in creased or decreased in proportion to the size of the pollen grains; if large pollen grains or spores--diameter about 80 st or morc, are to be cut sometimes as few as 1-3 pollen grains will be sufficient, cf. Fig. 258). 3. the concentration of the pollen grains (cf. Fig. 259; if the concentration > LHS STEEL HOLDER Fig 237. Preliminary cutting of the plastic unit.
>POLLEN GRAINS »OBJECT-HOLDER Fig. 258. Plastic unit with ready-made pyramid. (NB The Pyramid is enlarged about 10 times, the rest of the figure about 3 times ) is low, the dimensions of the pyramid, with a view to increasing the number of pollen grains to be sectioned, must be larger). The total height of the pyramid should not exceed half a millimetre if sections of a thickness of about 0 25-0 50 u are to be made) These recommendations have been followed when working with the modified Spencer microtome model 821 and are not always valid for ultra-thin sectioning. The best results are obtained by working with samples rich in pollen grains and absolutely clean. Organic debris renders cutting more difficult, shreds the sectioned pollen fragments, or covers them, and decreases the value of the sections on the whole. Inorganic material (e g. silicon or other mineral particles) will damage the microtome knife. Fig. 260 illustrates the order, directions etc., to be followed when moulding the pyramid, but it goes without saying that there can be many modı- PLASTIC SECTIONED POLLEN Fig 259 Seclion from the basal part of the pyramid. x 100. 10-505817 @ Endıman
140
OBJECT-HOLDER
B
INCISION NI,
PLASTIC UNIT
A
INC. No 2
B
INC. No 3
TING Nou
ING
No5
D
Fig. 260. Moulding of the pyramid. (N.B. Plastic unit 4:1, object-holder
about 2:1 ) - A. incision No. 1 - B. The object-hokler is turned 180°
and incisions Nos. 2 and 3 are made. A part of the area of the plastic unit
will fall away with incision No. 3, after which the object-holder is turned
again 180°, - C. Incision No. #cuts off the other part of the apes of the
plastic unit Two sides of the pyramid are now ready. The nhject-holder is
then turned 90°. D. Incision No. 5. D, shows the object-holder with
the plastic unit in top perspective after incision No. 5. - 1. The object
141 INC NA 6 E INC No 7 F INC No 8 G Gi holder is then turned 180° and incisions Nos 6 and 7 are made, after which it is again turned 180°, – F. Incision No 8 cuts away the last part of the plastic unit, and the coarse cutting of the pyramid is now complete. ---- G. Side view. G, Front view of the object-holder with ready-cut pyramid, fications according to individual ways of working, Usually, the moulding cannot be considered to be complete in eight cuts, and further fine trimming along the lines illustrated in Fig. 260 is always necessary. The height of a properly moulded pyramid is always greater than the base. The grinding of the microlome knife American “Eversharp" razor blades are used for cutting. After removing the fat with ether the blade is mounted in a special holder (LKB-Produkter, Stochliolm). The grinding is performed on a sheet of planed glass. A suitable amount of grinding powder (type B. 5125, Linde Air Products Co.) and
142
THE ARM OF THE MICROTOME
MOVES VERTICALLY. THE POINT OF THE PYRAMID
MUST BE PLACED EXACTLY ON THE
SAME LEVEL AS THE KNIFE.
TROUGH
KNIFE
LIQUID
INITIAL DISTANCE
OBJECT HOLDER
WITH PLASTIC
UNIT
Fig. 261 Adjusting of the Spencer microlome model 821 for sectioning.
teepol (teepol one part and distilled water four parts) should be available,
The powder is thoroughly mixed with the tecpol to a homogeneous, casya
flowing mixture. Ordinarily, grinding takes about 15 minutes. The pressure
on the knife should be at a minimum during the last five to eight minutes
of grinding. After rinsing in running water and drying with a few drops of
absolute alcohol, the knife is ready to use. It should be used inimediately
after grinding in order to avoid rust, contamination by dust etc.
The sectioning
The knife is cleansed with acetone and mounted in a curved position in
a special trough (made by LKB-Produkter, Stockholm). Care must be
143 MICROTOME THE ARM OF THE MICROTOME IS NOT IN MOTION AT THIS STAGE. MICROSCOPE TROUGH TERMINAL POSITION INITIAL POSITION > SHOOTING FORWARD l'ing 262 Atljusting of the metulome for sectiomng (rontinnet) taken to ensure that its edge is parallel with the end of the trough. The latter, together with the knife, is then fixed in the microtome. At the same time, the object-holder is tightly screwed into the mobile part of the microtome. The screw controlling the thich ness of the sections is adjusted in accordance with the desired thickness. Concomitantly, a final adjustment of the pyramid in relation to the knife should be made. The general conditions of statıhty must also be observed, particularly a steady undercarriage. This process, as well as that of cutting, is folloned through a binocular microscope (20).
IN THE FA ONT PERSPECTIVE THE POINT OF THE PYRAMID IS PLACED EXACTLY AT RIGHT ANGLES TO THE EDGE OF THE KNIFE OBJECT-HOLDER PLASTIC PYRAMID KNIFE TROUGH Fig 263. Pyramid and trough, front view. By hand, and with the motor off, the pyramid is brought to the level of the knife. In order to prevent damage to the pyramid through the vertical movement of the microtome, the trough with the kniſc should be placed at a safe distance. The knife is then slowly brought forward to the apex of the pyramid. Fig. 261 outlines the mobile part of the microtome, the knife-trough part (side view), and the vertical adjustment of the lever of the microtome with the mounted plastic unit. Fig. 262 shows the knife-trough block and the microscope. The final adjusting for sectioning is followed under the microscope. In the visual field will be seen the facet of the knife, the liquid (20 per cent alcohol in water) with which the trough has been filled, and the plastic unit with a part of its holder. One of the base lines of the pyramid must be parallel to the facet of the knife which should be brought as ncar as possible to the apex of the pyramid without coming into contact with it. The knife-trough block is then fixed, and the level of the liquid checked (the surface should be slightly convex). The motor is then started. Some introductory movements of the micro tome carry the apex of the pyramid towards the knifc. The microtome ſeeds at every second stroke. Normally, the sections assemble in the 143 SIDE VIEW THE PYRAMID IS ADJUSTED HORIZONTALLY KNIFE ty SURFACE OF THE LIQUID TROUGH AND STEEL PLATE IN CROSS SECTION STEEL PLATE Fig 264 Pyramid and part of trough, side view. trough, grouping themselves into wreaths, strips, or small flocks. If there is too much liquid they will disseminate and, furthermore, water will possibly adhere to the pyramid Inspection of the sections through the microscope will be rendered difficult or impossible by disturbing light refraction, resulting from an exaggerated convexity of the surface of the liquid etc. Occasionally, the sections will roll themselves up on the surface of the liquid, but they will regain their original shape iſ allowed to float in the trough for a few minutes before fishing up. -The above methods of embedding, cutting of pyramids, grinding and sectioning have been recommended by, la, F. S. Sjostrand, Karolinska Institutet, Stockholm. The fishing-up of the sections The sections are fished up from the trough with a thin, fine-pointed, soft brush, and transferred to a slide which has been thoroughly cicansed (albumin glycerine is not used) A drop of alcohol (20 per cent) is placed on the centre of the slide. The brush is then inserted into the liquid and slowly pushed towards that part of the knife where the sections are assembled. When the tip comes under the sections the brush is carefully lifted up over the surface of the water, turned upside down, and allowed to touch the surface of the liquid on the slide. At this stage a preliminary COVER-GLASS WITH SECTIONEO POLLEN FRAGMENTS EMBED- DED IN GLYCERINE- JELLY. LAVATERA PALLIDA SECTION THICKNESS 9,54 FUCHSIN 18mm PARAFINIZED AREA UNDER A COVER GLASS Fig. 265. Microscope slide with sectioned pollen walls. check under the microscope can be made in order to get an den of the quality of the sections. Further treatment depends on the number of sections transferred to the slide. If there are only a few it is better to allow the liquid on the slide to dry in the air. If the slide contains a great number of sections it is better to wash it with one or two drops of absolute alcohol. Although this procedure will probably wash away a number of the sections it is, nevertheless, worth while because the alcohol will separate the sections from each other and, at the same time, remove any dust etc. on the slide. When dry, the slide is rinsed once more with absolute alcohol. After the alcohol has evaporated the plastic is dissolved in acetone. The dissolving of the plastic, staining and mounting The acetone usually dissolves the plastic in a few hours (generally, tlie slide is left over-night in the acetone). After inspection vnder the microscope at low magnification (about <60) good section-bearing areas are marked with a diamond. The slide is then again carefully rinsed with absolute alcohol and with slowly running water. If the sections are not to be stained, the slide is ready for mounting. Staining with (lichsin, safranine, or methylene blue is useful, and in fact aften necessary, particularly if the thickness of the sections is only 147 0.5 pl or less. The stains are dissolved in cuvettes filled with water (approximately a knife's point to a cuvette). Staining is effected in about 4-5 hours. Bismarck brown, malachite green, and other stains have been tried, but with less satisfactory results. Staining greatly facilitates the localizing of the sections as unstained sections thinner then 0.3 p are often difficult to locate under an ordinary microscope. After staining, the slide is rinsed with water and allowed to dry Meanwhile, minute glycerine jelly cubes (about 1 mm) are cut, and a number of cover-slips carefully cleaned. The cubes are placed on those parts of the slide which have previously been marked with the diamond. The size of the cubes should vary in relation to the size of the section-bearing areas. A cover-slip is then placed on top of each jelly cube, and the jelly slowly melted. Care should be taken to prevent the jelly from spreading to the edge of the cover-slip. The slide is then sealed with paraffin wax. To this end a small piece of paraffın (melting point about 70 centigrades) is placed the edge of th cover-slip. The slide is then slowly heated in order to melt the paraffin which is sucked in under the cover-slip forming a protecting zone around the section-bearing glycerine jelly (Fig. 265). When the paraffın has cooled and hardened, the slide is cleaned with a knife and cotton wool soaked in benzol. For further information see "Sjostrand Ultra-Microtome. Instructions for use". (Distributed by LKB-Produkter, Stockholm 12.)
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INDEX The references to plastrations are distinguished by heavy rigures Generic names are i italics. The names of species are not included Abies 4, 6,7 Armopyle 7 Acroschurma 100 Acrostichum 46 Actinopteris 46 Acinostrobus & Adiantopsis 46 Adiantum 47 Agathis 8 Alnus 131 Alsophila 47 mentolarus 8 Amphidesmium 47 Amplectator 60 Anarthropleris 48 Andreaeaceae 99 Anemia 78 Aneuraceae 99 Angiopteridaceae 45 Angiopleris 79 Angectangulim 100 Anthoceros 100 Anthocerotaceae 99 Aposaccale 3 Aposaccia 3 Araucaria 9 Araucariaceae 5 Arreuthos 26 Irchangiopteris 79 Archidiaceae 99 Irchidium 101 Aspleniopsis 48 .1splenium 49 Athalamia 101 throtaxis 9 Athyrium 49 ustrotarus 9 Arolla 50 Azollaceae 45 Blasia 101 Blasiaceae 99 Blechnum 50 Blindla 103 Bulbutis 51 Botrychium 83 Bowenia 10 Brachiolejeunia 102 Brainca 51 Bryophyta 99 Bryaceae 99, 102 Bryum 102 Buxbaumia 102 Buxbaumiaceae 19 Buxbaumiales 102 Callris 10 Calomniaceae 99 Calomnium 103 Calymperaceae 99 Calymperes 103 Cappa 3 Cappula 3 Catascopiacrae 99 Calnscopium 112 Cedrus 4, 10, 11, PII (facing P 12) Cephalotaxaceae 5 Cephalolaru 12, PI I (lacing p. 12) Ceratodon 103 Cerafopteris 52, 53 Ceterach 53 Chamaecyparis 23, 26 Cherropleuria 54 Christensenin 79 Christenseniaceae 45 Cubotium 54 Cincidium 102 Clebea 104 Cleveaceae 93 Clivia 128 Cnemídaria 55 Cobaea 128 Codoniaceae 99 Conoceptialaccae 99 Conocephalum 104 Corpus 3 Corpus (lieight, breadthi, depth) 4 Corsinta 105 Corsiniaceae 99 Cristae marginales Cristae proximales 4 Cryptogramma 55 Cryptomeria 12 Cucurbila 128 Cunninghamia 12 Cupressaceac 5, 26 Cupressus 23, 24 Cyathea 56 Cyatheaccae 45 Cyathodiaceae 09 Cyathodrum 105 Cycadaceac 5 Cycas 13, 20 Cyclophorus 56 Cyrtomium 56 Cystodium 90 Cystopteris 56, 57 Dacryduum 13, 14-18 Danaca 79 Danaeaceae 45 Davallıq 57 Dendroceros 106 Dennstaedlta 57 Dichsonia 58 Dich soniaceae 45 Dicranaceae 99 Dicranales 103 Diffraction lines 127 Dioon 18 Diplıysciaccae 99 Diphyscium 102 Diplazium 58 Dipteridaceae 15 Dipteris 58 Diseima 18 Ditrichaceae 99 Drymoglossum 59 Drynatia 59 Dryoplcris 60 Dumortiera 106 Elaphoglossum 60 Encalypta 107 Encalyptaceae 99 Ephedra 4, 19, 20; Pl. 11 (facing p. 20) Ephedraceae 5 Ephemeraceae 99 Eguisetaceae 45 Equisetum 60 Euosmunda 84 Erormotheca 108 Exormothecaccac 19 Fegatella 104 Fissidens 108 Fissidentaceae II Fitzroya 26 Fossombronia 108 Fragmentation 128 Frullania 109 Frullaniaceae 99 Funaria 129, 130, PP IV, V (facing pp. 130, 131) Funariaceae 93 Gaura 128 Georgia 109 Georgiaceae 99 Geothallus 109 Gigaspermaceae 99 Gigaspermum 110 Ginkgo 21 Ginkgoaceae 5 Glercheniaceae 45, 63 Glyptostrobus 21 Gnetaceae 5 Grelum 22 Grammits 61 Grimaldiaceae 39 Grimmiaceae 99 Gymnogramme 61 Gymnospermae 5 Gymnospermae, megaspores 5 Gymnospermae, pollen grains 5 IIedwigia 110 Hedwigiaceae 99, 110 Jedwigidram 110 Helicophyllaceae 09 Helicophyllum 110 Jlemtohitis 61 Hemilelia 62 Hepaticae 99 Ilieriopteris 63 Histiopteris 63 Boltlumiella 64 Hookeria 111 Hookeriaceae 99 Humala 64 Hylacomiaceae 19 Ilylocomeant 111 Ilymenoglossum 96 }gmenolepis 65 Hymenoplayllaceae 45, 96 Hymenophyllopsidaceae 45 Hymenophyllopsis 65, 66 Ilymenophyllum 66, 96 Ilypolepis 67 Interference microscopy 131 Isoetaceae 45 Isoeles 67, 68 Isaloma 69 Isothecium 113 Jamesania 69 Juniperus Frontispiece, 24-26 Keleleeria 27 Lamtum 128 Large 28 Lastrea 69 Lembophyllaceae 99 Leptolepia 70 Leptopteris 84 Leptogtornaceae 93 Leplostomum 111 Liskea 111 Lesbeaceae 99 Leucobryaceae 99 Leucobryum 103 Liboredrus 26 Lindsaea 70 Linum 128 Lophosoria 71, 72 Lorentziella 110 Lorinseria 73 Lorsoma 73 Lorsomaceae 15 Lorsomopsis 74 Lycopodiaceae 45 Lycopodium 74, 75-7 129, 131; P!. I11 (tacia p. 34) Lygodrum 78 Macroglena 96 Macroglossum 79 MQerosamia 29 Noratlia 79 Marattiaceae 45 Narattiales 79 Marchantiaceae 99 Marsilea 80 Marsileaceae 45 Matomaceae -15 Marleucia 80 Neesea 112 Meeseaceae 99, 112 Megaceros 112 Mesosaccale 3 Mesosaccia 3 Welasequoia 29 Methacrylate 133 Microcachrys 30 Microlepia 80 Micropuncta 3 Microstrobus 30 Niechhoferia 102 Mitochondria 130 Mnraccae 39, 102 Afohria 78 Vonoclea 113 Monocfeaceae 99 Musci 99 Nanomitrium 113 Neclera 113 Neckeraceae 99 Necherinales 113 Megripteris 81 Neocallilropar 30 Nephrolepis 81 Yothorurus 37 Oedipodiaceae 99 Oedipodim 114 Oenothera 128 Oleandra 81 noclea 82 Onychium 82 151 Ophioglossaceae 45 Ophioglossum 83 Optical path difference 127, 131 Ormoloma 84 Orlhtopleris 84 Orthotrichaceae 99 Osmic acid 126 Osmunda 84 Osmundaceae 45 Osmundastrum 84 Orymitru 114 Oxymitraceae 99 Paludella 112 Parkeriaceae 45 Pellaea 85 Pellia 115 Pelliaceae 99 Peranema 85 Phanerophlebia 85 Phanerosorus 86 Phascum 117 Pherosphaera 3, 30 Phlebodrum 86 Phyllocladus 31 Phylloglossum 87 Picea 32, 33 Pilgerodendron 34 Pilularta 87 Palulariaceae 45 Pinaceae 5 Pinus 34-36 Plagrochasma 115 Plagiochila 116 Plagiochilaceae 99 Plagrogyria 88 Plagiogyriaceae 15 Plastids 130 Platycerium 88 Plalyzoma 63 Pleasium 84 Pleurophascaceae 99 Pleurophascum 116 Podocarpaceac 5 Podocarpus 36, 37, 38 Pohlia 102 Polarization microscopy 131 Polybotryja 88 Polypodiaceae 45 Polytrichaceae 99 Polytrichum 116 Poltia 117 Pottiaceac 99, 117 Protocyatheaceae 45 Pseudolari: 39 Pseudotsuga 40 Psilotaceae 45 Psilotum 89 Pleridium 89 Pteridophyta 45 Pteridophyta, megaspores 45 Pteridophyta, isospores and microspores 45 Pyrrhobryum 117 Pyrrosia 89 Radula 118 Radulaceae 99 Replica methods 128 Rhacomtrium 118 Rhizogoniaceae 99 Riccardıq 118 Riccia 119 Ricciaceae 99 Riella 120 Riellaceae 99 Russula 128 Saccale 3 Sacci 3 Saccoloma 89 Saccus (height, breadth, depth) 4 Salvinia 90 Salviniaceae 45 Saregothaea 40 Scapanta 121 Scapaniaceae 99 Schistostega 121 Schistostegaceae 99 Schizaea 78 Schizaeaceae 45, 78 Sciadopitys 41 Selaginella 90-94, P! III (facing p 91) Selaginellaceae 45 Selenodesmium 96 Sequoia 41 Sequoiadendron 41 Serpyllopsis 72 Selosisporites 2 Southbya 121 Sphacrocarpaceae 99 Sphagnaccae 99 Sphagnum 121 Spiridens 122 Spiridentaceae 99 Splachnaceae 99 Splachrum 122 Staining 146 Stangeria 42 Stenochlaena 94 Sticherus 63 Stroemia 124 Stromatopteris 63 Symphyodon 122 Symphyodontaceae 99 Symphyogyna 123 Symphyogynaceae 09 Taiwanra 42 Taenitis 95 Tapeinidium 95 Targionia 123 Targioniaceae 99 Taxaceae 5 Taxodiaceae 5 Taxodum 42 Tarus 42 Tayloria 122 Tenuitas 3 Thujopsis 43 Thyrsopteris 95 Timmia 124 Timmiaceae 99 Tmesipteridaceac 45 Tmesipferis 96 Todea 84 Torreya 43 Trichomanes 96 Tsuga 3, 43, 44 Ulota 124 Vandenboschta 96 l'itlarla 96 Vortia 122 Welwitschra 4, 19 Welwitschiaccac 3 Widdringlonta 26 Blood'sta 97 Xiphopteris 97 X-ray diffraction 131
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