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74 THE PROKARYOTIC ALGAE

Fig. 2.59 Light micrographs of cyanobacteria in the Nostocales. (From Rascher et al., 2003.)

Fig. 2.60 Jindong Zhao Dr. Zhao received his Bachelor’s degree in biology from Southwestern Normal University in 1982. He became a Ph.D. student in the Department of Botany, University of Texas at Austin in 1984 and received a Ph.D. degree in Biology in 1989. He spent the next three years as a postdoctoral fellow in Penn State University and the following two years in Applied Biosystems Inc. as a research scientist. Jindong Zhao returned to China and became an associate professor in the College of Life Sciences of Peking University in 1994. He was promoted to full professor in 1998. His laboratory focuses on studies of molecular mechanism of heterocyst differentiation and pattern formation of cyanobacteria.

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16:5744–8.

Part III

Evolution of the chloroplast

Fig. III.1 Diagrammatic representation of the uptake of a cyanobacterium by a protozoan into a food vesicle. This resulted in the establishment of an endosymbiosis between the cyanobacterium and the protozoan. Through evolution, the endosymbiotic cyanobacterium evolved into a chloroplast surrounded by two membranes of the chloroplast envelope.

The Rhodophyta (red algae) and Chlorophyta (green algae) form a natural group of algae in that they have chloroplasts surrounded by only the two membranes of the chloroplast envelope. The evolutionary event that led to the chloroplast occurred as follows (Fig. III.1). A phagocytotic protozoan took up a cyanobacterium into a food vesicle. Instead of being digested as a source of food, the cyanobacterium lived as an endosymbiont in the protozoan. This event benefited the protozoan because it received some of the photosynthate from the endosymbiotic alga, and it benefited the cyanobacterium because it received a protected stable environment. Through evolution the wall of the endosymbiotic

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cyanobacterium was lost. A mutation in the endosymbiont which resulted in a loss of the wall would have been selected for in evolution because it would have facilitated the transfer of compounds between the host and the endosymbiont. The food vesicle membrane of the phagocytotic host became the outer membrane of the chloroplast envelope. The plasma membrane of the cyanobacterium symbiont became the inner membrane of the chloroplast envelope. Rearrangement of the thylakoid membranes and evolution of polyhedral bodies into a pyrenoid completed the transition to a true chloroplast such as occurs in extant green and red algae.

The endosymbiotic origin of chloroplasts was first proposed by the Russian biologist Konstantin Mereschkowsky (1855–1921) (Fig. III.2) with the fundamentals of the idea appearing in his 1905 work, The Nature and Origins of Chromatophores in the Plant Kingdom (see Martin and Kowallik, 1999, for English translation). Mereschkowsky worked on the symbiosis between algae and fungi in lichens around the city of Kazan and this led him to his theory on endosymbiosis. He likened plastids to “little green slaves” working for their host cells to produce food from sunshine.

Subsequent cytological and biochemical studies have reinforced Mereschkowsky’s theory of an endosymbiotic origin of plastids. Nucleotide sequencing (either directly from rRNA or from DNA encoding rRNAs) has shown the similarity between cyanobacteria and plastids. It has further shown that all plastids evolved from a single endosymbiotic event (McFadden, 2001; Keeling, 2004).

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Fig. III.2 Konstantin Mereschkowsky Born August 4, 1855 in Warsaw, Poland (then part of Russia). Died by suicide in the Hotel des Familles, Geneva, Switzerland, on January 9, 1921. His father was a high official in the court of the Czar. He was the eldest son, having five brothers and three sisters. He entered the University of St. Petersburg in 1875, where he graduated with the diploma of Kandidat (with distinction) in 1880. In 1883 he became a Privatdocent at the same university. In 1883 he married Sultanova Olga Petronina by whom he had a son, Boris. From 1881 to 1898 he worked as a pomologist on fruit trees in the Crimea. In 1898 he left his wife and son to a life of poverty and traveled to the United States where he did research on unicellular algae (which he started when he was in the Crimea) at the University of California, Berkeley.

In 1902 he returned to Russia and took the position of curator of the zoology museum at Kazan University. In 1903 he passed his Magister with a dissertation “On the Morphology of Diatoms.” At that time Mereschkowsky’s research was influenced by Andrei Famintsyn, at the University of St. Petersburg, who was trying to cultivate chloroplasts isolated from zooxanthellae. Famintsyn was, however, not interested in having a younger competitor and denigrated Mereschkowsky’s research, resulting in bad relations between the two. In 1905 Mereschkowsky published his famous paper in Russian and German “On the Nature and Origin of Chromatophores in the Plant Kingdom.” In 1920, in Geneva, he published his most extensive paper on

the endosymbiotic origin of chloroplasts “La plant considérée comme un complexe symbiotique.”

Mereschkowsky’s personal life was a disaster. As a student he had revolutionary ideas. However, by the turn of the century he was a collaborator of the Czar’s secret police. He was one of the organizers of the nationalistic, anti-Semitic organization “The Kazan Department of the Union of Russian People.” Several of his colleagues were discharged after his denunciations. Mereschkowsky was forced to leave Russia in 1914, not because of his political views, but because of a sex scandal involving pedophilic activities between 1905 and 1914 with 26 young girls. The public scandal was nationwide. He fled to France in February 1918 and finally took refuge in Switzerland. Near the end of his life he lamented that his theory on the symbiotic origin of chloroplast “had made little headway” and that it was “often completely ignored.” This was compounded by “finally the war, the revolution.” After a particularly odious exchange with R. Chodat, Professor of Botany at the University of Geneva, who prevented Mereschkowsky from speaking, Mereschkowsky commented “One day history will remember me.” On Sunday, January 9, 1921, having run out of money, and after paying all his hotel bills, he committed suicide by gassing himself with chloroform. Thus passed away at 65 years of age the most famous evolutionary botanist of the century. For a well written and researched biography of Mereschkowsky see Sapp et al. (2002).

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