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Floridean starch

Floridean starch is a type of a storage glucan found in glaucophytes and in red


algae (also known as rhodophytes), in which it is usually the primary sink for fixed
carbon from photosynthesis. It is found in grains or granules in the cell's cytoplasm
and is composed of an α-linked glucose polymer with a degree of branching
intermediate between amylopectin and glycogen, though more similar to the former.
The polymers that make up floridean starch are sometimes referred to as "semi-
amylopectin".[1]

Illustration of glucose polymer


Contents branching.

Properties
Evolution
History
References

Properties
Floridean starch consists of a polymer of glucose molecules connected primarily by α(1,4) linkages, with occasional branch points
using α(1,6) linkages. It differs from other common α-linked glucose polymers in the frequency and position of the branches, which
gives rise to different physical properties. The structure of floridean starch polymers is most similar to amylopectin and is sometimes
described as "semi-amylopectin". Floridean starch is often described in contrast to
starch (a mixture of amylopectin and amylose) and
glycogen:[1]

Floridean starch Starch Glycogen


Some bacteria, some
Organisms Red algae, glaucophytes Green algae, plants
archaea, fungi, animals
Semi-amylopectin; classically without
Composition amylose, though some examples exist Amylopectin and amylose Glycogen
with amylose present
Storage
In the cytosol Inside plastids In the cytosol
location
Eukaryotes: UDP-glucose
Building
UDP-glucose ADP-glucose Bacteria: ADP-glucose
block

Amylopectin: Branches are


relatively rare and occur in
clusters Branches are relatively
Branching Intermediate level of branching frequent and evenly
Amylose: Almost entirely
distributed
linear

Genes
required for Fewer than 12 30–40 6–12
maintenance
Historically, floridean starch has been described as lacking amylose. However, amylose has been identified as a component of
[2][3]
floridean starch granules in some cases, particularly in unicellular red algae.

Evolution
Features such as UDP-glucose building blocks and cytosolic storage differentiate the Archaeplastida into two groups: the
rhodophytes and glaucophytes, which use floridean starch, and the green algae and plants (Chloroplastida), which use amylopectin
and amylose. There is strong phylogenomic evidence that the Archaeplastida are monophyletic and originate from a single primary
endosymbiosis event involving a heterotrophic eukaryote and a photosynthetic cyanobacterium.[1][4]

Evidence indicates that both ancestors would have had established mechanisms for carbon storage. Based on review of the genetic
complement of modern plastid genomes, the last common ancestor of the Archaeplastida is hypothesized to have possessed a
cytosolic storage mechanism and to have lost most of the endosymbiotic cyanobacterium's corresponding genes.[1][5] According to
this hypothesis, the rhodophytes and glaucophytes retained the ancestral eukaryote's cytosolic starch deposition. Starch synthesis and
degradation in green algae and plants is much more complex – but significantly, many of the enzymes that perform these metabolic
[1][2]
functions in the interior of modern plastids are identifiably of eukaryotic rather than bacterial origin.

In a few cases, red algae have been found to use cytosolic glycogen rather than floridean starch as a storage polymer; examples such
as Galdieria sulphuraria are found in the Cyanidiales, which are unicellularextremophiles.[6][7]

Other organisms whose evolutionary history suggests secondary endosymbiosis of a red alga also use storage polymers similar to
floridean starch, for example, dinoflagellates and cryptophytes. The presence of floridean starch-like storage in some apicomplexan
parasites is one piece of evidence supporting a red alga ancestry for theapicoplast, a non-photosynthetic organelle.[8]

History
Floridean starch is named for a class of red algae, the Florideae (now usually termed Florideophyceae).[9] It was first identified in the
mid-19th century and extensively studied bybiochemists in the mid-20th century.[10]

References
1. Ball, S.; Colleoni, C.; Cenci, U.; Raj, J. N.; T
irtiaux, C. (10 January 2011). "The evolution of glycogen and starch
metabolism in eukaryotes gives molecular clues to understand the establishment of plastid endosymbiosis". Journal
of Experimental Botany. 62 (6): 1775–1801. doi:10.1093/jxb/erq411 (https://doi.org/10.1093%2Fjxb%2Ferq411).
PMID 21220783 (https://www.ncbi.nlm.nih.gov/pubmed/21220783).
2. Ball, Stephen; Colleoni, Christophe; Arias, Maria Cecilia (2015). "The ransition
T from Glycogen to Starch Metabolism
in Cyanobacteria and Eukaryotes". In Nakamura, Y asunori. Starch: Metabolism and Structure(https://link.springer.co
m/chapter/10.1007/978-4-431-55495-0_4). Springer Japan. pp. 93–158.ISBN 978-4-431-55494-3.
3. McCracken, D. A.; Cain, J. R. (May 1981). "Amylose in Floridean Starch".
New Phytologist. 88 (1): 67–71.
doi:10.1111/j.1469-8137.1981.tb04568.x(https://doi.org/10.1111%2Fj.1469-8137.1981.tb04568.x) .
4. Viola, R.; Nyvall, P.; Pedersen, M. (7 July 2001). "The unique features of starch metabolism in red algae"(https://ww
w.ncbi.nlm.nih.gov/pmc/articles/PMC1088757). Proceedings of the Royal Society B: Biological Sciences . 268 (1474):
1417–1422. doi:10.1098/rspb.2001.1644(https://doi.org/10.1098%2Frspb.2001.1644). PMC 1088757 (https://www.n
cbi.nlm.nih.gov/pmc/articles/PMC1088757) . PMID 11429143 (https://www.ncbi.nlm.nih.gov/pubmed/11429143).
5. Dauvillée, David; Deschamps, Philippe; Ral, Jean-Philippe; Plancke, Charlotte; Putaux, Jean-Luc; Devassine, Jimi;
Durand-Terrasson, Amandine; Devin, Aline; Ball, Steven G. (15 December 2009)."Genetic dissection of floridean
starch synthesis in the cytosol of the model dinoflagellate"(https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2795531).
Proceedings of the National Academy of Sciences . 106 (50): 21126–21130. doi:10.1073/pnas.0907424106(https://d
oi.org/10.1073%2Fpnas.0907424106). PMC 2795531 (https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2795531) .
PMID 19940244 (https://www.ncbi.nlm.nih.gov/pubmed/19940244).
6. Martinez-Garcia, Marta; Stuart, Marc C.A.; van der Maarel, Marc J.E.C (August 2016). "Characterization of the highly
branched glycogen from the thermoacidophilic red microalga Galdieria sulphuraria and comparison with other
glycogens". International Journal of Biological Macromolecules
. 89: 12–18. doi:10.1016/j.ijbiomac.2016.04.051(http
s://doi.org/10.1016%2Fj.ijbiomac.2016.04.051) .
7. Deschamps, Philippe; Haferkamp, Ilka; d’Hulst, Christophe; Neuhaus, H. Ekkehard; Ball, Steven G. (November
2008). "The relocation of starch metabolism to chloroplasts: when, why and how".Trends in Plant Science. 13 (11):
574–582. doi:10.1016/j.tplants.2008.08.009(https://doi.org/10.1016%2Fj.tplants.2008.08.009).
8. Coppin, Alexandra; Varré, Jean-Stéphane; Lienard, Luc; Dauvillée, David; Guérardel, Y ann; Soyer-Gobillard, Marie-
Odile; Buléon, Alain; Ball, Steven; T
omavo, Stanislas (February 2005). "Evolution of Plant-Like Crystalline Storage
Polysaccharide in the Protozoan Parasite Toxoplasma gondii Argues for a Red Alga Ancestry".Journal of Molecular
Evolution. 60 (2): 257–267. doi:10.1007/s00239-004-0185-6(https://doi.org/10.1007%2Fs00239-004-0185-6) .
PMID 15785854 (https://www.ncbi.nlm.nih.gov/pubmed/15785854).
9. Barry, V. C.; Halsall, T. G.; Hirst, E. L.; Jones,J. K. N. (1949). "313. The polysaccharides of the florideœ. Floridean
starch". Journal of the Chemical Society: 1468–1470. doi:10.1039/JR9490001468(https://doi.org/10.1039%2FJR949
0001468).
10. Meeuse, B. J. D.; Andries, M.; Wood, J. A. (1960). "Floridean Starch".Journal of Experimental Botany. 11 (2): 129–
140. doi:10.1093/jxb/11.2.129 (https://doi.org/10.1093%2Fjxb%2F11.2.129) .

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