BSCI 411

PLANT GENETICS and MOLECULAR BIOLOGY

Department of Cell Biology & Molecular Genetics


Organelle Genomes

I. Non-Mendelian Inheritance
	-maternal inheritence- depends on parent

II. Origin- Endosymbiotic Theory     1-2 billion yrs ago
	mitochondria- purple photosynthetic bacteria
	plastid-chloroplast- cyanobacteria
	similarities:
		circular DNA
		DNA lacks histones
		ribosomes similar 70S (50 and 30)
		inhibited by similar antibiotics (e.g chloramphenicol, streptomycin)
		different GC content ( 36%) than nuclear (64%)
Intermediates in Endosymbiosis-secondary endosymbiosis (only with plastid)
	-in certain algae-chlorarachinophytes ameboid algae
		see two sets of rDNA-chloroplast has 4 membranes
		genome outside of chloroplast is nucleomorph
	others- another algae which swallowed red algae
			-Phylum Apicomplexa
		-plasmodium (malarial parasite), toxiplasma
			-plastid but non-photosynthetic- secondary plastid from green algae
	
III. Chloroplast Genome
-22-900 copies per plastid
	-associated in aggregate called nucleoids
-Overall organization
	-120-400 kB in size 
	-large inverted repeat (IR) 
	-3 classes, based upon presence of inverted repeat
-Genes
	-100 protein coding genes, rDNA, tRNA
	-loss of 90% of cyanobacteria genes  (3,500 genes)
	genes in chloroplast
		a. ~40 Photosynthetic genes
		b. ~ 50 needed for replication, transcrition, translation
                        rRNA, some tRNA, polymerase
		c. unknown
			-variable  e.g. rice & maize have only 1
			-red algae 30 others- aa synthesis, protein transport

IV.  Mitochondrial Genome
	sizes 208(watermelon) -2,500 kB (muskmelon)
-Comparison with animals
		-human- simplest 16,569 bp , compact
			-tightly packed  2 rDNA
					22tRNA
					5 membrane proteins
					8 unassigned
			-2 RNA transcripts make copies of each half
				-tRNAs flank mRNAs- all cut from two transcripts during 
                                       transcription
		-yeast
			-5x as much DNA10 open reading frames and 6 membrane proteins
			genes separated by AT rich stretches

-plants- 5x larger than yeast
			- genes in mt- more genes than animals (Bryophyte 94 ORFs)
				rRNA 26S, 18S, 5S (plants only)
                                ribosomal proteins- one in some plant mt 
				tRNA- full complement
				protein-coding
					some of inner membrane complexe’s subunits
					-replication, transcription, translation
					cytochrome C
					ATPase
                                           (mammals- subunit 9 in nucleus, plants, 
                                             yeast in mt)
					apocytochrome B of bc1 complex

	-Organization- tripartite(unlike animals- single circle)
		-master circle and smaller subgenomic circles	
			-master produces two genomics through recombination
                Why not a mess of fragments?		
  		    Recombination gives produces variability, mutation rate 10x lower than nucleus
  		    -e.g cytoplasmic male sterility in maize
	              -mt gene- tURF13  114 codons
		         88 aa similar to rrn26 (ribosomal RNA)
		          8 unknown
		         18 more similar to rrn26
		         flanking 5 ‘ region(5kB) atp26 gene
                      -in anthers, some anther-specific gene interacts- makes mt leaky so dies
		      -T-toxin in Cochliobolus maydis does same
			
	-RNA editing:
	      -discrepancies between DNA sequence and protein sequence in wheat
		-originally thought to be deviation in code CGG arg->trp
		-sequenced cDNA found cGG changed to UGG
	            -C ->> U, sometimes U->> C
		-depends on gene, no consensus

	    -other editing
               -mt and cht already unusual in cutting RNA to give multiple subtranscripts of 
                       mRNA as well as rRNA and tRNA (latter two normal)
               - cis and trans splicing, often combined
		
Promiscuous genes- found in different compartments
	-Cyanophora- rbcS still in chloroplast, next to rbcL
	-chloroplast sequences in mt DNA of maize
	-Many of these look like reverse-transcribed copies
	-introns also hopping around- self splicing type I introns found to move, also see 
	     movement from fungi into plants



Links

Organelle genome database
Example of secondary endosymbiosis
Chloroplast genome organization- PLSC731 NDSU
Mitochondrial genome organization- PLSC731 NDSU
Summary of Mitochondrial Genome
RNA editing- PLSC731 NDSU

Last revised: March 17 1998- Straney