John F. Allen. Cell Biology and Developmental Genetics. Lectures by. jfallen.org
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2 Cell Biology and Developmental Genetics Lectures by John F. Allen School of Biological and Chemical Sciences, Queen Mary, University of London jfallen.org 1
3 Cell Biology and Developmental Genetics Lectures by John F. Allen
4 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history
5 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history Endosymbiosis and the origin of bioenergetic organelles. A modern view
6 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history Endosymbiosis and the origin of bioenergetic organelles. A modern view Mitochondria as we know them and don't know them
7 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history Endosymbiosis and the origin of bioenergetic organelles. A modern view Mitochondria as we know them and don't know them Why do chloroplasts and mitochondria have genomes?
8 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history Endosymbiosis and the origin of bioenergetic organelles. A modern view Mitochondria as we know them and don't know them Why do chloroplasts and mitochondria have genomes? Co-location for Redox Regulation
9 Cell Biology and Developmental Genetics Lectures by John F. Allen Endosymbiosis and the origin of bioenergetic organelles. Some history Endosymbiosis and the origin of bioenergetic organelles. A modern view Mitochondria as we know them and don't know them Why do chloroplasts and mitochondria have genomes? Co-location for Redox Regulation Mitochondria, ageing, and sex energy versus fidelity
10 Cell Biology and Developmental Genetics Lectures by John F. Allen
11 Cell Biology and Developmental Genetics Lectures by John F. Allen Slides and supplementary information:
12 Cell Biology and Developmental Genetics Lectures by John F. Allen Slides and supplementary information: jfallen.org/lectures
13 Lecture 1 Endosymbiosis and the origin of bioenergetic organelles. Some history
14 What are bioenergetic organelles?
15 What are bioenergetic Chloroplasts organelles?
16 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells
17 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis
18 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression
19 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression
20 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression Mitochondria
21 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression Mitochondria Membrane-bound compartments in eukaryotic cells
22 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression Mitochondria Membrane-bound compartments in eukaryotic cells Function in energy conversion in respiration
23 What are bioenergetic organelles? Chloroplasts Membrane-bound compartments in eukaryotic cells Function in energy conversion in photosynthesis Contain a genome and a complete apparatus of gene expression Mitochondria Membrane-bound compartments in eukaryotic cells Function in energy conversion in respiration Contain a genome and a complete apparatus of gene expression
24
25 Figure Electron micrographs of chloroplasts. (A) In a wheat leaf cell, a thin rim of cytoplasm containing chloroplasts, the nucleus, and mitochondria surrounds a large vacuole. (B) A thin section of a single chloroplast, showing the chloroplast envelope, starch granules, and lipid (fat) droplets that have accumulated in the stroma as a result of the biosyntheses occurring there. (C) A high-magnification view of two grana. A granum is a stack of thylakoids. (Courtesy of K. Plaskitt.) 2002 by Bruce Alberts, Alexander Johnson, Julian Lewis, Martin Raff, Keith Roberts, and Peter Walter.
26 Figure The chloroplast. This photosynthetic organelle contains three distinct membranes (the outer membrane, the inner membrane, and the thylakoid membrane) that define three separate internal compartments (the intermembrane space, the stroma, and the thylakoid space). The thylakoid membrane contains all the energy-generating systems of the chloroplast, including its chlorophyll. In electron micrographs, this membrane seems to be broken up into separate units that enclose individual flattened vesicles (see Figure 14-34), but these are probably joined into a single, highly folded membrane in each chloroplast. As indicated, the individual thylakoids are interconnected, and they tend to stack to form grana by Bruce Alberts, Alexander Johnson, Julian Lewis, Martin Raff, Keith Roberts, and Peter Walter.
27 Figure A mitochondrion and chloroplast compared. A chloroplast is generally much larger than a mitochondrion and contains, in addition to an outer and inner membrane, a thylakoid membrane enclosing a thylakoid space. Unlike the chloroplast inner membrane, the inner mitochondrial membrane is folded into cristae to increase its surface area by Bruce Alberts, Alexander Johnson, Julian Lewis, Martin Raff, Keith Roberts, and Peter Walter.
28 What is symbiosis? Simon Schwendener (1867) discovered that lichens are a consortium of two organisms - a fungus and an alga Heinrich Anton de Bary (1878) used symbiosis ( same living ) to describe this association
29 The origin of the Endosymbiont hypothesis
30 Konstantin Sergejewicz Mereschkowsky ( ) Constantin Mereschkowsky, Über Natur und Ursprung der Chromatophoren im Pflanzenreiche. Biol. Centralbl., 25 (1905): English translation by Martin, W. And Kowallik, K. (1999) Eur. J. Phycol. 34:
31 Background to the endosymbiont hypothesis, 1: plastids in Arabidopsis thaliana plastids multiply by division as observed repeatedly in plants and algae in 19 th century
32 Background to the endosymbiont hypothesis, 2: cyanobacteria plastids Small, blue-green, round or oval bodies of very simple structure Small, green (probably blue-green originally, as with Cyanomonas), round or oval bodies of very simple structure The green pigment saturates the plasm uniformly or is distributed therein in the form of small droplets The green pigment saturates the stroma uniformly or is distributed therein in the form of small droplets
33 Background to the endosymbiont hypothesis, 3: cyanobacteria plastids Do not possess a true nucleus, rather just certain structures (nucleic granules) that can be viewed as a predecessor of the nucleus Do not possess a true nucleus, rather certain structures (pyrenoids) that can be viewed as a transformed, primitive nucleus Nutrition: assimilation of CO 2 in the light Reproduction: through division Nutrition: assimilation of CO 2 in the light Reproduction: through division
34 Plastids are organs
35 Plastids are organs accepted view was that plastids were organs
36 Plastids are organs accepted view was that plastids were organs autonomously derived from cytoplasm
37 Plastids are organs accepted view was that plastids were organs autonomously derived from cytoplasm Mereschkowsky: this notion is, by no means, a finding supported by direct observation
38 Plastids are organs soybean
39 Plastids are organs soybean why was it thought plastids arose de novo?
40 Plastids are organs soybean why was it thought plastids arose de novo? plant parts that are kept in the dark turn green when exposed to light
41 Plastids are organs soybean why was it thought plastids arose de novo? plant parts that are kept in the dark turn green when exposed to light Schimper (1885) showed that chlorophyll arises de novo but not its carriers
42 Plastids are not organs
43 Plastids are not organs plastids do not arise de novo but from pre-existing (sometimes colourless) plastids
44 Plastids are not organs plastids do not arise de novo but from pre-existing (sometimes colourless) plastids Mereschkowsky: this hypothesis is...founded upon an error in logic. And those who wish to maintain their view...reveal that they lack a proper grasp of the nature of heredity
45 Continuity of plastids
46 Continuity of plastids plastids arise from pre-existing plastids, never de novo
47 Continuity of plastids plastids arise from pre-existing plastids, never de novo so, at one stage, a plastid established itself in a plant lineage
48 Continuity of plastids plastids arise from pre-existing plastids, never de novo so, at one stage, a plastid established itself in a plant lineage therefore, plastids must be foreign bodies or symbionts
49 Plastids are independent
50 nucleus is central to cell Plastids are independent
51 Plastids are independent nucleus is central to cell if nucleus is removed, cells do not proliferate
52 Plastids are independent nucleus is central to cell if nucleus is removed, cells do not proliferate plastids continue to function in enucleated cells
53 Plastids are independent nucleus is central to cell if nucleus is removed, cells do not proliferate plastids continue to function in enucleated cells grow and multiply by division
54 Plastids are independent nucleus is central to cell if nucleus is removed, cells do not proliferate plastids continue to function in enucleated cells grow and multiply by division assimilate CO 2
55 Plastids are independent
56 Plastids are independent plastids function biochemically even in vitro
57 Plastids are independent plastids function biochemically even in vitro plastid produce lipids that are distinct from those from the rest of the cell
58 Plastids are independent plastids function biochemically even in vitro plastid produce lipids that are distinct from those from the rest of the cell plastids have a colourless mass (pyrenoid) similar to bacteria (this is their DNA but unknown at the time) and produce protein
59 Plastids are independent plastids function biochemically even in vitro plastid produce lipids that are distinct from those from the rest of the cell plastids have a colourless mass (pyrenoid) similar to bacteria (this is their DNA but unknown at the time) and produce protein so, plastids behave like independent organisms
60 polyp Hydra viridis contains endosymbiotic zoochlorellae (algae) Analogy with hydra
61 Analogy with hydra polyp Hydra viridis contains endosymbiotic zoochlorellae (algae) some hydra contain symbionts that behave as plastids in plants
62 Analogy with hydra polyp Hydra viridis contains endosymbiotic zoochlorellae (algae) some hydra contain symbionts that behave as plastids in plants these also originate by division
63 Analogy with hydra polyp Hydra viridis contains endosymbiotic zoochlorellae (algae) some hydra contain symbionts that behave as plastids in plants these also originate by division these symbionts can live outside the host
64 Free-living plastids..? Mereschkowsky: A theory such as the one suggested here would gain considerably...if...free-living organisms with similarity to the symbionts could be demonstrated cyanobacteria (cyanophytes; blue-green algae)
65 Anabaena cylindrica Watercolour, 1946, by G. E. Fogg
66 Endosymbiosis with cyanobacteria cyanobacterium, Richelia intercellularis in a diatom, Rhizosolenia styliformis rhizopod Paulinella chromatophora parasitized by cyanobacteria Living examples are known
67 Plastids are endosymbionts
68 Plastids are endosymbionts plastids arise by division from previous plastids
69 Plastids are endosymbionts plastids arise by division from previous plastids plastids continue to function normally in enucleated cells
70 Plastids are endosymbionts plastids arise by division from previous plastids plastids continue to function normally in enucleated cells plastids make protein
71 Plastids are endosymbionts plastids arise by division from previous plastids plastids continue to function normally in enucleated cells plastids make protein cyanobacteria seem very similar to plastids
72 Plastids are endosymbionts plastids arise by division from previous plastids plastids continue to function normally in enucleated cells plastids make protein cyanobacteria seem very similar to plastids cyanobacteria are known that live in symbioses with other organisms
73 Further consequences
74 Further consequences According to this theory, the plant cell is nothing other than an animal cell with cyanobacteria that have invaded it
75 Further consequences According to this theory, the plant cell is nothing other than an animal cell with cyanobacteria that have invaded it the green, the brown and the red algae could have thus originated independently
76 Further consequences According to this theory, the plant cell is nothing other than an animal cell with cyanobacteria that have invaded it the green, the brown and the red algae could have thus originated independently All of the peculiarities that are characteristic for plants and that distinguish them from animals, are the natural result of a symbiosis between an animal cell and a CO 2 - assimilating cyanobacterium
77 Discredited until the 1970s
78 Discredited until the 1970s Popular theory in early 20 th century
79 Discredited until the 1970s Popular theory in early 20 th century Wallin postulated a similar theory for mitochondria in 1927
80 Discredited until the 1970s Popular theory in early 20 th century Wallin postulated a similar theory for mitochondria in 1927 Wilson ridiculed these ideas: To many, no doubt, such speculations may appear too fantastic for present mention in polite biological society...
81 Discredited until the 1970s Popular theory in early 20 th century Wallin postulated a similar theory for mitochondria in 1927 Wilson ridiculed these ideas: To many, no doubt, such speculations may appear too fantastic for present mention in polite biological society... These hypotheses were made before molecular biology, before it was known that DNA was the hereditary material
82 Typical prokaryotic (left) and eukaryotic (right) cells. W. Ford Doolittle Nature 392, 15-16, 1998
83 The endosymbiont hypothesis for the origin of mitochondria. W. Ford Doolittle Nature 392, 15-16, 1998
84 Summary
85 Summary Mereschkowsky proposed 100 years ago that the bodies - chromatophores found in green plants and algae are enslaved cyanobacteria
86 Summary Mereschkowsky proposed 100 years ago that the bodies - chromatophores found in green plants and algae are enslaved cyanobacteria The endosymbiont hypothesis for the origin of chloroplasts and mitochondria is now orthodox, but there is controversy over the nature of the symbionts and their place in eukaryote evolution
87 The origin of the Endosymbiont hypothesis
88 The origin of the Endosymbiont hypothesis Konstantin Sergejewiz Merezkovskij ( )
89 The origin of the Endosymbiont hypothesis Konstantin Sergejewiz Merezkovskij ( ) (University in Kasan, Russia)
90 The origin of the Endosymbiont hypothesis Konstantin Sergejewiz Merezkovskij ( ) (University in Kasan, Russia) Constantin Mereschkowsky, Über Natur und Ursprung der Chromatophoren im Pflanzenreiche. Biol. Centralbl., 25 (1905):
91 The origin of the Endosymbiont hypothesis Konstantin Sergejewiz Merezkovskij ( ) (University in Kasan, Russia) Constantin Mereschkowsky, Über Natur und Ursprung der Chromatophoren im Pflanzenreiche. Biol. Centralbl., 25 (1905): Constantin Mereschkowsky, Über Natur und Ursprung der Chromatophoren im Pflanzenreiche. Biol. Centralbl., 25 (1905): English translation by Martin, W. And Kowallik, K. (1999) Eur. J. Phycol. 34:
92 Konstantin Sergejewicz Mereschkowsky ( )
93 The renaissance of the Endosymbiont hypothesis
94 The renaissance of the Endosymbiont hypothesis Lynn Margulis (b. 1938)
95 The renaissance of the Endosymbiont hypothesis Lynn Margulis (b. 1938) (University of Massachussetts, Amhurst)
96 The renaissance of the Endosymbiont hypothesis Lynn Margulis (b. 1938) (University of Massachussetts, Amhurst) Margulis, Lynn, 1970, Origin of Eukaryotic Cells, Yale University Press
97 The renaissance of the Endosymbiont hypothesis Lynn Margulis (b. 1938) (University of Massachussetts, Amhurst) Margulis, Lynn, 1970, Origin of Eukaryotic Cells, Yale University Press Margulis, Lynn, 1992, Symbiosis in Cell Evolution: Microbial Communities in the Archean and Proterozoic Eons, W.H. Freeman
98 Lynn Margulis (b. 1938)
99 Lecture 2. October 28th, a.m. Endosymbiosis and the origin of bioenergetic organelles. A modern view
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