Classification of uniserial striae in Bacillariophyta with bipolar frustule
The striae are the main structural-functional units in the diatom frustule, however, their morphology is still insufficiently studied and poorly used in Bacillariophyta taxonomy. New criteria for the stria architecture analysis rely on the data from electron microscopy investigations. For the first...
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Цитувати: | Classification of uniserial striae in Bacillariophyta with bipolar frustule / L.N. Bukhtiyarova // Альгология. — 2015. — Т. 25, № 2. — С. 198-210. — Бібліогр.: 19 назв. — англ. |
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irk-123456789-1099552016-12-25T03:02:43Z Classification of uniserial striae in Bacillariophyta with bipolar frustule Bukhtiyarova, L.N. Систематика, филогения и вопросы эволюции водорослей The striae are the main structural-functional units in the diatom frustule, however, their morphology is still insufficiently studied and poorly used in Bacillariophyta taxonomy. New criteria for the stria architecture analysis rely on the data from electron microscopy investigations. For the first time hierarchical classification of the striae diversity in the diatom bipolar frustules was proposed. It reflects species ontogeny and the influence of ecology in evolution of different striae. Штрихи являются основными структурно-функциональными единицами панциря Bacillariophyta, однако их морфология остается не достаточно изученной и ограничено используется в таксономии Bacillariophyta. Новые критерии хтя анализа архитектуры штриха базируются на данных электронной микроскопии. Впервые предложена иерархическая классификация разнообразия штрихов у Bacillariophyta с биполярной симметрией панциря. Она отражает онтогенез видов и влияние экологии на эволюцию разнообразных штрихов. 2015 Article Classification of uniserial striae in Bacillariophyta with bipolar frustule / L.N. Bukhtiyarova // Альгология. — 2015. — Т. 25, № 2. — С. 198-210. — Бібліогр.: 19 назв. — англ. 0868-8540 DOI: http://doi.org/10.15407/alg25.02.198 http://dspace.nbuv.gov.ua/handle/123456789/109955 582.26: 556.55 en Альгология Інститут ботаніки ім. М.Г. Холодного НАН України |
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Систематика, филогения и вопросы эволюции водорослей Систематика, филогения и вопросы эволюции водорослей |
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Систематика, филогения и вопросы эволюции водорослей Систематика, филогения и вопросы эволюции водорослей Bukhtiyarova, L.N. Classification of uniserial striae in Bacillariophyta with bipolar frustule Альгология |
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The striae are the main structural-functional units in the diatom frustule, however, their morphology is still insufficiently studied and poorly used in Bacillariophyta taxonomy. New criteria for the stria architecture analysis rely on the data from electron microscopy investigations. For the first time hierarchical classification of the striae diversity in the diatom bipolar frustules was proposed. It reflects species ontogeny and the influence of ecology in evolution of different striae. |
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Bukhtiyarova, L.N. |
author_facet |
Bukhtiyarova, L.N. |
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Bukhtiyarova, L.N. |
title |
Classification of uniserial striae in Bacillariophyta with bipolar frustule |
title_short |
Classification of uniserial striae in Bacillariophyta with bipolar frustule |
title_full |
Classification of uniserial striae in Bacillariophyta with bipolar frustule |
title_fullStr |
Classification of uniserial striae in Bacillariophyta with bipolar frustule |
title_full_unstemmed |
Classification of uniserial striae in Bacillariophyta with bipolar frustule |
title_sort |
classification of uniserial striae in bacillariophyta with bipolar frustule |
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Інститут ботаніки ім. М.Г. Холодного НАН України |
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2015 |
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Систематика, филогения и вопросы эволюции водорослей |
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http://dspace.nbuv.gov.ua/handle/123456789/109955 |
citation_txt |
Classification of uniserial striae in Bacillariophyta with bipolar frustule / L.N. Bukhtiyarova // Альгология. — 2015. — Т. 25, № 2. — С. 198-210. — Бібліогр.: 19 назв. — англ. |
series |
Альгология |
work_keys_str_mv |
AT bukhtiyarovaln classificationofuniserialstriaeinbacillariophytawithbipolarfrustule |
first_indexed |
2025-07-07T23:53:22Z |
last_indexed |
2025-07-07T23:53:22Z |
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1837034271994281984 |
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198
ISSN 0868-8540. lgologia. 2015, 25(2): 198—210 http://dx.doi.org/10.15407/alg25.02.198
UDK 582.26: 556.55
L.N. BUKHTIYAROVA
Institute for Evolutionary Ecology, NAS of Ukraine,
Akad. Lebedev St., 37, Kiev 03143, Ukraine
CLASSIFICATION OF UNISERIAL STRIAE IN BACILLARIOPHYTA
WITH BIPOLAR FRUSTULE
The striae are the main structural-functional units in the diatom frustule, however, their
morphology is still insufficiently studied and poorly used in Bacillariophyta taxonomy. New
criteria for the stria architecture analysis rely on the data from electron microscopy
investigations. For the first time hierarchical classification of the striae diversity in the
diatom bipolar frustules was proposed. It reflects species ontogeny and the influence of
ecology in evolution of different striae. Striae types were determined according to areolae
types and originally defined stria parameters. Morphology of areolae determines
macroareolar, postmacroareolar, combined and polyareolar striae types which are considered
as the consistent stages in evolution of the striae in bipolar frustule. Our concept on the
derivation of prevail in Bacillariophyta polyareolar stria from the macroareolar stria through
the intermediate stria types is supported by similar morphological stages and their
consistency recorded in frustule ontogeny of the species from different systematic groups of
Bacillariophyta with bipolar symmetry. Basal and interbasal stria types were defined in
accordance with the stria location within a basal siliceous layer. Conical type of areola and
polyareolar stria with conical areolae are described. Their distinction from poroid areolae and
polyareolar stria with poroid areolae is discussed. Proportional, compact and distant kinds of
striae are defined on the ratio between stria and interstria areas and their meaning in species
biology and ecology is indicated. All the definitions of the frustule structural units are based
on the concept of functional morphology of the diatom frustule suggested by the author.
K e y w o r d s : functional morphology, db-element, df-morph, terminology, macroareola,
postmacroareola, poroid areola, conical areola; stria: basal, interbasal, macroareolar,
postmacroareolar, combined, polyareolar, proportional, distant, compact, transformed;
frustule: monosymmetrical, bisymmetrical, three-symmetrical.
Introduction
For a long time the number of the rows with areolae within a stria was the
only character of stria morphology applied in description of the diatom taxa.
Diatoms studied with electron microscopy displayed large diversity in stria
morphology. However, these data are not yet reflected in modern systematic
of Bacillariophyta. At present only alveolate or chambered stria type is
distinguished in the diatom frustule. An alveola was defined as ‘‘an elongated
© L.N. Bukhtiyarova, 2015
Classification of uniserial striae
199
chamber or one of a series of elongated chambers running from the axial or
central part of the valve to the margin, opening to the inside of the frustule by
a large opening and with an areolate outer layer’’, for example, in Pinnularia
viridis (Nitzsch) Ehrenb. and species of Cyclotella Kütz. (Ross et al., 1979:
p. 527). Later the term alveolate stria was suggested (Cox, Ross, 1981).
Different kinds of “chambering” in raphid genera Progonoia H.-J. Schrad.,
Scoliotropis Cleve, some species of Gomphonema Ehrenb. and Gomphoneis
Cleve, araphid Gephyria media Arn. were discussed from ontogenetic point of
view (Cox, 2012).
The aim of present study is to designate key morphological parameters of
stria that allow to distinguish them precisely and to propose the definitions for
the striae diversity found in different Bacillariophyta taxa with bipolar frustule.
Materials and methods
The materials for our study were collected over a long period in different
hydrotopes in the Ukraine. The micrographs of Psammothidium vernadskyi
Bukht. et Stanislav. were taken from the sample collected by I.L. Kusin from
little lake (Blue Lake N 29), Russia, East Siberia, 70 km south-east of city
Hanty-Mansiysk. Samples were treated on standard methods (Krammer,
Lange-Bertalot, 1986). Equipment includes scanning electron microscope
JEOL 6060LA, Japan; transmission electron microscope JEOL JEM 1230.
Results and discussion
General definitions
Frustule symmetry has significant importance in Bacillariophyta taxonomy.
Traditionally diatom frustule symmetry is defined on the quantity of
symmetric sections and axes that can cross it (Diatomovy …, 1949). Bilateral
bipolar frustule may possess following mutually perpendicular planes of
symmetry: valvar ( in Ukr., in
Rus.), longitudinal ( in Ukr., in
Rus.) and transverse ones ( in Ukr.,
in Rus.). Different bilateral bipolar frustules can be crossed on one—three
pairs of equal (symmetrical) parts. Thus, the term bilateral is insufficient for
the precise description of the diatom frustule symmetry. Therefore we have
introduced following new terms.
Monosymmetrical frustule (Eng., in Ukr.,
in Rus.) — frustule that can be crossed with one
plane of symmetry. This type of frustule symmetry characterizes of
Rhoicosphaenia Grunow (symmetrical on longitudinal plane of symmetry).
Bisymmetrical frustule (Eng., in Ukr., -
in Rus.) — frustule that can be crossed with two planes of
symmetry. An example of such frustule symmetry is Eunotia (symmetrical on
valvar and transversal planes of symmetry).
Three-symmetrical frustule (Eng., in Ukr.,
in Rus.) — frustule that can be crossed with three
L.N. Bukhtiyarova
200
planes of symmetry. An example of this frustule symmetry is Navicula Bory
(symmetrical on valvar, longitudinal and transversal planes of symmetry).
For the first time proposed terminology was applied in the descriptions of
the new taxa from Lake Baikal (Bukhtiyarova and Pomazkina, 2013).
For the species with bilateral bipolar symmetry a stria were redefined as
“… depressions sunk into surface of the valve” (Cox and Ross, 1981). In the
terms of functional morphology stria of the diatom frustule (
in Ukr., in
Rus.) is a universal repeated micro-df-morph of third (second) order, the area
with areolae restricted by neighboring virgae. Stria functions provide
interaction of protoplast with environment through the hierarchical system:
df-converter—areola—stria. Different processes, stria functions, take place in
stria area: selective concentration of the matters on their surface, matters
absorption/excretion, intensification of physiological processes connected with
cell membrane, focusing and filtering of light and others. Evolution of a stria
reflects the development of its functions depending on particular
environmental conditions (Bukhtiyarova, 2009).
Basic element of the diatom frustule (= db-element, Eng.,
, = db- in Ukr.,
, = db- in Rus.) —
morphologically detached, homogeneous frustule component, that possesses
special physical-chemical features and provides a primary base for the diatom
frustule hierarchical construction. They belong to db-elements of a diatom
frustule as different apertures and cavities in its thickness, regularly repeated
and unique silica components.
Morph of the diatom frustule (= df-morph, Eng.;
, df- in Ukr.;
, df- in Rus.) — compound structural unit of the diatom
frustule that consists the db-elements or structural units of lower orders, or
both of them, realizes particular functions for the diatom organism and has
own evolution (Bukhtiyarova, 2009).
The term virga was proposed as the “… bars between the striae” to
distinguish interstrial bars into interior of the frustule in pinnate diatoms of
Naviculaceae or Eunotiaceae from the true costae in Diatomaceae or some
Nitzschiaceae (Cox and Ross, 1981). In the same paper the term vimin was
suggested for the “crossbars” between virgae. Lately virga was defined
“… ontogenetically as the transverse bars that develop from the sternum”
(Cox, 2012). Our definitions are based on functional morphology.
Virga (Lat., in Ukr., in Rus.) — universal repeated siliceous
micro-db-element in form of long siliceous bar that restricts stria area and
protects it from mechanical stress (Bukhtiyarova 2009: figs 1—6: v).
Vimin (Lat., in Ukr., in Rus.) — universal repeated
siliceous nano-db-element in form of the plates or short bars between areolae.
The type and arrangement of the vimines within striae area determine the
pattern of areolae arrangement in a stria (Bukhtiyarova, 2009: figs 2—5: n).
Classification of uniserial striae
201
Areola (Lat., in Ukr., in Rus.) — universal repeated nano-
(micro-) df-morph of third/second order that includes a hollow in the frustule
siliceous layer, one or two df-converters and organized in a stria. Functions of
areola coincide partially with the stria functions. It is obvious that df-converter
morphology defines the most important characteristics of areola. However,
other parameters, like areola hollow length and shape into the frustule
thickness, its inner surface etc, determine areola functions as well
(Bukhtiyarova, 2009).
onverter of the diatom frustule (=df-converter, Eng., df- in
Ukr., df- in Rus.) syn.: velum, occlusion — universal, repeated
nano- (micro-) df-morph of first order that consists fine aperture (-s) and
siliceous db-element (-s), locates within areola and carries out direct
interaction between protoplast and environment: matters absorption/
excretions, light energy focusing and filtering; biological, chemical protecting
and others. Df-converter transforms the effects of environmental factors in
suitable for diatom organism manner and values however these basic functions
are not reflected in any of earlier suggested terms which only indicate the fact
of areola closing (Bukhtiyarova, 2009).
Types of striae
In our opinion, the type of stria is determined by the type of areolae,
their arrangement within stria area and other stria parameters. Morphology of
areolae assigns following stria types.
Macroareolar stria (Lat., in Ukr., -
in Rus.) (Figs 3—5) — type of stria that consists of
macroareola.
According to definition, macroareola ( in Ukr.,
in Rus.) is a type of areola in which the foramen occupies the whole stria area
and closed by flat sieve df-converter (Bukhtiyarova, 2006: figs 3, 4). Thus,
macroareolar stria is equal morphologically to one macroareola.
Postmacroareolar stria (Lat., in Ukr.,
in Rus.) Figs 6—7a — type of stria that consists
of postmacroareolae. Several postmacroareolae may occupy a whole stria area.
Postmacroareola (Lat., in Ukr., in
Rus.) Figs 7a, 17, df-c — areola with large foramen of irregular size and shape
even within one stria, occupying large part of stria area and closed by sieve df-
converter (Bukhtiyarova, 2006: figs 1, 2).
Morphology of postmacroareolar stria clearly proves that in evolution
process it apparently was an intermediate stria type between macroareolar and
polyareolar stria, one of the prevalent stria types of different systematic groups
Bacillariphyta. In particularly, both in macroareolar stria and postmacroareolar
one the previmins often are present.
Previmin (Lat., in Ukr., in Rus.) fig. 7, arrows —
repeated siliceous nano-db-element in form of incomplete transverse plates of
irregular form and location partially crossed macroareola or postmacroareola
L.N. Bukhtiyarova
202
foramen. Their actual function is in reinforcing of striae area with large df-
convertes. The previmines evolutionally precede to the vimines.
Observations of stria development during valve ontogeny in the diatoms
with bipolar symmetry have showed that the vimins start to form when virgae
shaping has been completed and that became the base for a concept of stria as
a single areola covered by velum with transverse ribs (Schmid and Schulz,
1979). Such a pattern of valve morphogenesis was confirmed in different
systematic groups of Bacillariophyta with bipolar symmetry (Mayama and
Kobayasi, 1989; Cox, 2012). The result of comparative morphological study of
Karayevia species coincides with the ontogeny data therefore hypothesis about
derivation of stria with poroid areolae from macroareolar striae via
postmacroareolar striae was suggested (Bukhtiyarova, 2006). Recent data also
pointed out the appearance of the vimins on the late stages of virgae
formation (Cox, 2012: figs 5b, f, h) which morphology is similar to the
previmins on this ontogeny stage but differ from them in the regular location
within stria area. Thus, the presence of the previmins in macroareolae and
postmacroareolae indicates on similarity of evolutional processes of those
areola types and maintains our concept of polyareolar stria derivation.
Polyareolar stria (Lat., in Ukr.,
in Rus.) — type of stria that consists the groups of the areolae with
regular morphology and certain pattern of allocation within stria area (Figs 1,
10—12).
Stria with poroid areolae (= poroid stria Lat.-Eng.;
, in Ukr., ,
= in Rus.) — polyareolar type of stria that consists poroid
areolae in a row with uniform vimins (Figs 1, 10—12).
Poroid areola (Lat., in Ukr., in Rus.) —
type of areola with uniform tube-like hollow into the frustule silica, sieve df-
converter on outer and flap-like df-converter on inner valve surface (Fig. 1).
1 2
Figs 1, 2. Scheme section across the striae and areolae in the diatom frustule with bipolar
symmetry. 1 — Interbasal stria with poroid areolae; 2 — basal stria with conical areolae.
df-c — df-converter, f — outer areola foramen, f-c — flap-like df-converter, his — height of
interstria area, hs — height of stria area, wis — width of interstria area, ws — width of stria
area
Classification of uniserial striae
203
Poroid areola was initially described as an areola which hollow has not
noticeable widening into the valve thickness (Nikolaev, 1984).
Stria with conical areolae (= conic-areolar stria Lat.-Eng.;
, - in Ukr., c
, - in Rus.) — polyareolar
type of stria that consists conical areolae in a row with uniform vimins (Figs
13—15).
Conical areola (Lat., in Ukr., in Rus.) —
type of areola with truncated conical hollow into the valve thickness, usually,
small outer foramen and considerably larger inner foramen with sieve df-
converter (Figs 2, 13—15). Normally df-converters are set on the props or on
the tapes.
The key morphological differences of conical areola from poroid one are:
1) conical hollow into frustule silica instead tube-like one, 2) single
df-converter of sieve type on inner valve surface while poroid areola has both
inner sieve df-converter and outer flap-like df-converter of various
morphology (= vola accoding to Ross and Sims, 1972); Mann, 1981; =
foricula accoding to Cox, 2004). It was earlier underlined that “ … slit-like
external areola openings within the Naviculales (Table 2) are not described as
volate” Cox (2004). However, in our opinion the areolae in both the genera
Navicula Bory and Cymbella C. Agardh possess outer flap-like df-converter
with slit-like opening. An example of conical areolae in proportional striae
can be found on illustrations of Placoneis sovereignae (Hust.) Torgan et
Donadel (Torgan et al., 2010: figs 12, 14).
reola prop (Eng., c in Ukr., in Rus.) —
repeated nano-db-element in form of siliceous strut that serves for
df-converter fixing (Figs 12, 20—22, arrows).
The term prop or supporting prop was suggested earlier without any
definition (Cox, 2004).
reola tape (Eng., in Ukr., in Rus.) —
repeated nano-db-element in form of thin siliceous strip that serves for
df-converter fixing (Figs 19—22, arrows).
It is unknown yet exactly whether in the same valve both props and tapes
can be present (Figs 19—22) or in this case the false props are appeared as a
result of tape destroying. We suppose as well that tapes can be formed
between props and mask the later ones.
Combined stria (Eng., in Ukr.,
in Rus.) — polyareolar kind of stria that consists regular row (-s) of
areolae with uniform vimins and postmacroareolae.
Combined striae are relatively rare in the diatoms with bipolar frustule but
occur in different genera and systematic orders that indirectly prove their
intermediate position in stria evolution. Usually postmacroareolae in
combined striae are allocated at valve margin or/and at valve mantle:
Psammothidium frigidum (Hust.) Bukht. et Round (Bukhtiyarova and Round,
1996: fig. 58), Fistulifera saprophyla (Lange-Bert. et Bonik) Lange-Bert.
L.N. Bukhtiyarova
204
(Lange-Bertalot, 2001: Pl. 111, fig. 4), Fistulifera pelliculosa (Bréb.) Lange-
Bert. (Lange-Bertalot, 2001: Pl. 112, fig. 1).
Thus, combined striae can be considered as second intermediate stage in
derivation of polyareolar stria from macroareolar one through postmacro-
areolar and combined striae.
Parameters of stria in the diatom frustule
Besides of morphologically different areolae following parameters are
characterized the striae of the diatom frustule.
Stria area (Sst) (Eng., in Ukr., in Rus.) —
the diatom frustule area restricted by neighboring virgae that a set of areolae
occupies.
Interstria area (Sist) (Eng., in Ukr.,
in Rus.) — the diatom frustule area that virga occupies.
The term interstria in fact was not accepted in today diatom morphology
and recently has been distinguished from the virgae: “… interstriae recognized
under LM may not correspond to virgae sensu Cox and Ross (1981). Virgae
are defined ontogenetically as the transverse bars that develop from the
sternum in pennate diatoms” (Cox, 2012: p. 18). However, as it was shown
above, the ontogeny pattern of the striae in bipolar frustule is similar in
taxonomically different diatoms both in general stages and their sequence.
Therefore the term virga has universal and more wide meaning than in it’s
original definition. Moreover, virgae have subordinate, assistant functions
regarding to the striae — the main functional unit in the diatom frustule
(Bukhtiyarova, 2009). So, according to our concept of stria architecture there
is no need to discuss virgae as the costae or not costae etc., since all the
definitions focus attention on the stria morphology which includes virgae
morphology as a character of particular stria type. Besides that the proposed
here term interstria area is not a morphological structure but one of a frustule
parameter.
Interstria height (Hist) (Eng.,
in Ukr., in Rus.) —
thickness of the basal siliceous layer in virga area (Figs 1, 2)
Depending on stria parameters following stria types were observed in the
diatom frustules with bipolar symmetry.
Basal stria (Eng., in Ukr., in Rus.) —
type of stria that areolae and all additional db-elements accompanied them are
sited within interstria height (Figs 2, 10—15). This stria type species of
Achnanthidium Kütz., Placoneis Mereschk. have.
Interbasal stria (Eng., in Ukr.,
in Rus.) — type of stria that areolae and additional db-elements
accompanied them are sited within a part of interstria height (Figs 1, 16—18).
This stria type the species of Navicula s. str. and Cymbella s. str. have.
Classification of uniserial striae
205
Kinds of striae
Proportional stria (Eng., in Ukr.,
in Rus.) — kind of stria that stria area is approximately equal to
interstria area (Sist : Sst = 1) (Figs 5, 7, 18).
Packed stria (Eng., in Ukr., in Rus.) —
kind of stria that stria area is two or more times as much than interstria area
(Sist : Sst 0.5) (Figs 3, 16).
Distant stria (Eng., in Ukr., in
Rus.) — kind of stria that stria area is two or more times as less than interstria
area (Sist : Sst 2) (Figs 4, 6, 8, 9, 10—13, 17).
Because the density of areolae on the valve with proportional and packed
striae are much higher than in the ones with distant striae it is obviously that
in species with first two kinds of striae the exchange of matters with
environment is much intensive than in species with distant striae. So, observed
gradation of striae morphology reflects biological characteristics of species and
can be useful in ecological investigations.
Variety of striae
Transformed stria (Eng., in Ukr., -
, in Rus.) — stria that consists the groups
of areolae with the foramens or the flap openings of the df-converters in
different size and transitional shape. Poroid striae with outer slit-like flap df-
converters often gradually changes the shape of the flap openings from slit-like
to shorter slits and even roundish ones especially at central valve part as in
many Navicula and Cymbella species (Fig. 17). In species with bi-
multyseriate striae the number of the rows with areolae may change gradually,
making a transformed striae like in Planothidium Round et Bukht.,
Gomphoneis Cleve. Variety of stria usually has species rank of taxonomy if
other species characters completely correspond to a genus diagnosis. In case
when the stria variety make up a set with other characters that distinguishes a
group of the species, it may correspond to genus rank of taxonomy.
Conclusions
Morphological criteria and essential principles of striae architecture in the
diatom frustule are grounded on the data of electron microscopy and
functional morphology. The proposed classification of newly described striae
includes co-subordinate units — types, kinds and variety in diversity of the
striae, which were found in the diatom species with bipolar symmetry. The
concept of an evolutional derivation of polyareolar striae from macroareolar
striae through the intermediate postmacroareolar and combined ones was
confirmed both with comparative morphological and ontogeny studies. The
progress in development of terminology will contribute to the exact and short
taxa diagnosis that is an actual task in today Bacillariophyta taxonomy.
The author greatly thanks to Dr. Nikolay Scherbatyuk for assistence on
taking of TEM micrographs, Dr. Sci. professor Michael Guiry, Ryan Institute,
Ireland, for English correction.
L.N. Bukhtiyarova
206
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2010.
Received 08.04.2014
Submitted by S.P. Wasser
ISSN 0868-8540. lgologia. 2015, 25(2): 198—210 http://dx.doi.org/10.15407/alg25.02.198
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Classification of uniserial striae
207
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L.N. Bukhtiyarova
208
Figs 3—9. Types of striae in the diarom frustule with bipolar symmetry. 3 — macroareolar striae,
Karayevia sp., fragment of raphe valve, outer surface; 4, 5 — macroareolar striae, interrupted at
valve mantle junction: 4 — distant striae, Karayevia sp. 1, fragment of rapheless valve, outer
surface, 5 — proportional striae, Amphora sp., outer surface, df-convertors are destroyed; 6, 7,
7a — postmacroareolar striae: 6 — distant striae, Brachysira sp., 7, 7a — proportional striae with
sieve df-convertors, Brachysira sp. 1; previmins, arrows; 8, 9 — basal, distant, transformed striae
with poroid areolae: 8 — fragment of valve Fragilaria sp., 9 — transformed striae, arrows, central
part of valve Fragilaria sp. 1, outer surface. a — areola, df-c — sieve df-converter, ma —
macroareola, pma — postmacroareola, n — vimin, v — virga. Figs 3—6, 9 — SEM; 7, 7a, 8 —
TEM. Scale 3, 5, 6, 9 — 1 m, 4 — 2 m, 7 — 250 nm, 7a — 100 nm, 8 — 200 nm
Classification of uniserial striae
209
Figs 10—18. Types of striae in the diarom frustule with bipolar symmetry. 10, 11 — polyareolar
basal distant striae with poroid areolae, Cocconeis sp., central part of raphe valve from outer and
inner surface; 12—15 — polyareolar basal striae with conical areolae: 12 — distant striae,
Encyonema sp., central part of valve from inner surface; areola props, arrows, 13 — distant striae,
Psammothidium sp., 14, 15 — proportional sriae, Psammothidium vernadskyi Bukht. et Stanislav.,
fragments of raphe valve from outer and inner surface; 16—18 — polyareolar interbasal striae with
poroid areolae: 16 — packed striae, Navicula sp., fragment from outer valve surface, 17 —
transformed striae in central part of Cymbella sp., arrows, outer valve surface, 18 — pro-
portional striae, Navicula sp. 1, fragment of inner valve surface. df-c — sieve df-converter,
n — vimin, v — virga. 10—12, 14—18 — SEM; 13 — TEM. Scale 10—12, 16—18 — 1 m,
14, 15 — 500 nm, 13 — 100 nm
L.N. Bukhtiyarova
210
Figs 18—21. reola db-elements and df-morphs, SEM. 18 — sieve df-converters in
compressed conical areolae, in light rectangles; stigma, arrow; fragment of Placogeia
microcephala Pomazk. et Bukht., outer valve surface; 19 — areola props, arrow; areola tape,
double arrow; fragment of Placogeia acceptaformis Bukht. et Pomazk., inner valve surface;
20 — areola tape, white arrow; props, black arrows; the pole of Baikalia aleksandrae
Pomazk. et Bukht., inner valve surface; 21 — areola tapes, white arrows; prop, black arrow;
additional sieve catinus, double arrows; the pole of Nadiya pavla Pomazk. et Bukht., inner
valve surface. Scale 19—22 — 1 m
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