Morphology
Visible structures
Two specimens of Laminaria hyperborea, each showing the rootlike holdfast at lower left, a divided blade at upper right, and a
stemlike stipe connecting the blade to the holdfast.
1 MORPHOLOGY
2.1
Fossils
3
lar ancestors.[18] DNA sequence comparison also suggests that the brown algae evolved from the lamentous Phaeothamniophyceae,[19] Xanthophyceae,[20] or the
Chrysophyceae[21] between 150[1] and 200 million years
ago.[2] In many ways, the evolution of the brown algae parallels that of the green algae and red algae,[22]
as all three groups possess complex multicellular species
with an alternation of generations. Analysis of 5S rRNA
sequences reveals much smaller evolutionary distances
among genera of the brown algae than among genera of
red or green algae,[2][23] which suggests that the brown
algae have diversied much more recently than the other
two groups.
2.1 Fossils
cur anywhere on the thallus.[11]
1.3
Tissue organization
The cell wall consists of two layers; the inner layer bears
the strength, and consists of cellulose; the outer wall layer
is mainly algin, and is gummy when wet but becomes hard Fossils comparable in morphology to brown algae are
known from strata as old as the Upper Ordovician,[29]
and brittle when it dries out.[15]
but the taxonomic anity of these impression fossils is
far from certain.[30] Claims that earlier Ediacaran fossils
are brown algae[31] have since been dismissed.[19] While
2 Evolutionary history
many carbonaceous fossils have been described from the
Precambrian, they are typically preserved as attened
Genetic and ultrastructural evidence place the Phaeo- outlines or fragments measuring only millimeters long.[32]
phyceae among the heterokonts (Stramenopiles),[16] Because these fossils lack features diagnostic for idena large assemblage of organisms that includes both tication at even the highest level, they are assigned to
photosynthetic members with plastids (such as the fossil form taxa according to their shape and other gross
diatoms) as well as non-photosynthetic groups (such as morphological features.[33] A number of Devonian fosthe slime nets and water molds). Although some het- sils termed fucoids, from their resemblance in outline
erokont relatives of the brown algae lack plastids in their to species in the genus Fucus, have proven to be inorcells, scientists believe this is a result of evolutionary loss ganic rather than true fossils.[24] The Devonian megafosof that organelle in those groups rather than independent sil Prototaxites, which consists of masses of laments
acquisition by the several photosynthetic members.[17] grouped into trunk-like axes, has been considered a possiThus, all heterokonts are believed to descend from a sin- ble brown alga.[11] However, modern research favors reingle heterotrophic ancestor that became photosynthetic terpretation of this fossil as a terrestrial fungus or fungalwhen it acquired plastids through endosymbiosis of an- like organism.[34] Likewise, the fossil Protosalvinia was
other unicellular eukaryote.[4]
once considered a possible brown alga, but is now thought
[35]
The closest relatives of the brown algae include unicel- to be an early land plant.
lular and lamentous species, but no unicellular species A number of Paleozoic fossils have been tentatively
of brown algae are known. However, most scientists classied with the brown algae, although most have
assume that the Phaeophyceae evolved from unicellu- also been compared to known red algae species.
Phascolophyllaphycus possesses numerous elongate, inated blades attached to a stipe. It is the most abundant of algal fossils found in a collection made from
Carboniferous strata in Illinois.[36] Each hollow blade
bears up to eight pneumatocysts at its base, and the stipes
appear to have been hollow and inated as well. This
combination of characteristics is similar to certain modern genera in the order Laminariales (kelps). Several fossils of Drydenia and a single specimen of Hungerfordia
from the Upper Devonian of New York have also been
compared to both brown and red algae.[26] Fossils of
Drydenia consist of an elliptical blade attached to a
branching lamentous holdfast, not unlike some species
of Laminaria, Porphyra, or Gigartina. The single known
specimen of Hungerfordia branches dichotomously into
lobes and resembles genera like Chondrus and Fucus[26]
or Dictyota.[37]
The earliest known fossils that can be assigned reliably
to the Phaeophyceae come from Miocene diatomite deposits of the Monterey Formation in California.[4] Several
soft-bodied brown macroalgae, such as Julescraneia, have
been found.[38]
CLASSIFICATION
Classication
3.1
Family
Phylogeny
1922
Family
Sphacelariaceae
Decaisne
1842
Taxonomy
Oltmanns
3.2
Stypocaulaceae
1922
the
orders
in
the
class
1925
Family
Desmarestiaceae
(Thuret)
Kjellman 1880
5
Family Phaeosiphoniellaceae Phillips
et al. 2008
[Chordariopsidaceae]
Family Scytosiphonaceae Ardissone
& Straforello 1877 [Chnoosporaceae
Setchell & Gardner 1925]
Family Petrospongiaceae Racault et al.
2009
4 Life cycle
Sexual reproduction may be isogamous, oogamous, or
anisogamous. There is evidence of sex chromosomes.[41]
Union of gametes may take place in water or within the
oogonium (oogamous species). The life cycle shows great
variability from one group to another.
In Laminaria, there is a conspicuous diploid generation.
Meiosis takes place within sporangia, before the spores
are released. As they are haploid, with sex chromosomes,
there are equal numbers of male and female spores.[42]
With the exception of the Fucales, all brown algae have a
life cycle with an alternation between haploid and diploid
forms.
Family Platysiphonaceae
Order Tilopteridales Bessey 1907 emend.
Phillips et al. 2008 [Cutleriales Bessey
1907]
5 Ecology
Family ?Masonophycaceae
Family Tilopteridaceae Kjellman 1890 Brown algae have adapted to a wide variety of marine
Family Phyllariaceae Tilden 1935
ecological niches including the tidal splash zone, rock
REFERENCES
8 See also
Wrack (seaweed)
9 References
[1] Medlin, L. K.; et al. (1997). Phylogenetic relationships
of the 'golden algae' (haptophytes, heterokont chromophytes) and their plastids (PDF). Plant Systematics and
Evolution. Plant Systematics and Evolution. 11: 187
219. doi:10.1007/978-3-7091-6542-3_11. ISBN 978-3211-83035-2. hdl:10013/epic.12690.
Chemistry
[12] Raven, P. H.; Evert, R. F.; Eichhorn, S. E. (2005). Biology of Plants (7th ed.). New York: W. H. Freeman and
Company. pp. 316321, 347. ISBN 0-7167-1007-2.
[13] Round, F. E. (1981). The Ecology of Algae. Cambridge:
Cambridge University Press. p. 103. ISBN 0-521-269067.
[15] Sharma, O. P (1986). Textbook of Algae. Tata McGrawHill. p. 298. ISBN 978-0-07-451928-8.
[16] Adl, S. M.; et al. (2005). The new higher level classication of eukaryotes with emphasis on the taxonomy of
protists (PDF). Journal of Eukaryotic Microbiology. 52
(5): 399451. doi:10.1111/j.1550-7408.2005.00053.x.
PMID 16248873.
[31] Loeblich, A. R. (1974). Protistan Phylogeny as Indicated by the Fossil Record. Taxon. 23 (2/3): 277290.
doi:10.2307/1218707. JSTOR 1218707.
[17] Lane, C. E.; Archibald, J. M. (2008). The eukaryotic tree of life: Endosymbiosis takes its TOL (PDF).
Trends in Ecology and Evolution. 23 (5): 268275.
doi:10.1016/j.tree.2008.02.004. PMID 18378040.
[18] Niklas, K. J. (1997). The Evolutionary Biology of Plants.
Chicago: University of Chicago Press. p. 156. ISBN 0226-58082-2.
[19] Lee, R. E. (2008). Phycology (4th ed.). Cambridge University Press. ISBN 978-0-521-63883-8.
[20] Ariztia, E. V.; Andersen, R. A.; Sogin, M. L.
(1991). A new phylogeny of chromophyte algae using 16S-like rRNA sequences from Mallomonas papillosa (Synurophyceae) and Tribonema aequale (Xanthophyceae)". Journal of Phycology. 27 (3): 428436.
doi:10.1111/j.0022-3646.1991.00428.x.
[21] Taylor, T. N.; Taylor, E. L. (1993). The Biology and Evolution of Fossil Plants. Englewood Clis, NJ: Prentice
Hall. pp. 128131. ISBN 0-13-651589-4.
[22] Dittmer, H. J. (1964). Phylogeny and Form in the Plant
Kingdom. Princeton, NJ: D. Van Nostrand Company. pp.
115137. ISBN 0-88275-167-0.
[23] Hori, H.; Osawa, S. (1987). Origin and evolution of organisms as deduced from 5S ribosomal RNS sequences
(PDF). Molecular Biology and Evolution. 4 (5): 445472.
PMID 2452957.
[24] Arnold, C. A. (1947). An Introduction to Paleobotany.
New York; London: McGraw-Hill. p. 48. ISBN 1-40671861-0.
[25] Coyer, J. A.; Smith, G. J.; Andersen, R. A. (2001).
Evolution of Macrocystis spp. (Phaeophyta) as determined by ITS1 and ITS2 sequences (PDF). Journal
of Phycology. 37 (4): 574585. doi:10.1046/j.15298817.2001.037001574.x.
[42] Thomas, D. N. (2002). Seaweeds. London: Natural History Museum, London. ISBN 0-565-09175-1.
[29] Fry, W. L. (1983). An algal ora from the Upper Ordovician of the Lake Winnipeg region, Manitoba, Canada.
Review of Palaeobotany and Palynology. 39 (34): 313
341. doi:10.1016/0034-6667(83)90018-0.
10
10
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11.2
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