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Where am I in the cell?

Control layer

Information

What the cell needs to know, where that knowledge is stored, and how it is read.

The genome

Aquifex aeolicus VF5 carries its genes on one chromosome of about 1.55 million base pairs and a small extra DNA element. UniProt lists 1,553 proteins encoded there. Aquifex is only the reference organism; the minimal cell has 1,051 proteins.

Source: Deckert et al., Nature 392:353–358 (1998)

Copying the genome

These proteins open, copy, join and untwist the DNA.

Aquifex proteinEvidence (UniProt)Home pathway
Chromosomal replication initiator protein DnaASeen as a proteinChromosome preparation
Replicative DNA helicaseSeen as a proteinInitiation and opening
DNA polymerase III subunit alphaInferred from related proteinsDNA synthesis
Beta sliding clampInferred from related proteinsDNA synthesis
DNA polymerase III subunit gamma/tauInferred from related proteinsDNA synthesis
Probable DNA polymerase III subunit deltaInferred from related proteinsDNA synthesis
DNA polymerase IInferred from related proteinsJoining and end processing
DNA ligaseInferred from related proteinsJoining and end processing
Type 2 topoisomerase subunit ASeen as a proteinTopology management
Type 2 topoisomerase subunit BSeen as a proteinTopology management
DNA topoisomerase 1Inferred from related proteinsTopology management
Single-stranded DNA-binding proteinInferred from related proteinsHomologous recombination
DNA primaseSeen as a proteinDNA replication

Reading genes

RNA polymerase copies genes into RNA. Only one sigma factor, which tells it where genes start, is kept.

Aquifex proteinEvidence (UniProt)Home pathway
DNA-directed RNA polymerase subunit alphaInferred from related proteinsTranscription
DNA-directed RNA polymerase subunit betaInferred from related proteinsTranscription
DNA-directed RNA polymerase subunit beta'Inferred from related proteinsTranscription
DNA-directed RNA polymerase subunit omegaInferred from related proteinsNot yet placed in a pathway
RNA polymerase sigma factor SigAInferred from related proteinsTranscription
Transcription termination/antitermination protein NusAInferred from related proteinsTranscription
Transcription antitermination protein NusBSeen as a proteinTranscription
Transcription termination/antitermination protein NusGSeen as a proteinTranscription

The code and its key

The genetic code links three-letter codons to amino acids. The key is a set of enzymes, the aminoacyl-tRNA synthetases, each of which attaches the right amino acid to its transfer RNAs. The kept set keeps 22 synthetase proteins (some enzymes are made of two different chains).

Aquifex proteinEvidence (UniProt)Home pathway
Alanine--tRNA ligaseSeen as a proteinTranslation interface
Arginine--tRNA ligaseInferred from related proteinsTranslation interface
Aspartate--tRNA(Asp/Asn) ligaseInferred from related proteinsTranslation interface
Cysteine--tRNA ligaseInferred from related proteinsTranslation interface
Glutamate--tRNA ligaseInferred from related proteinsTranslation interface
Glycine--tRNA ligase alpha subunitSeen as a proteinTranslation interface
Glycine--tRNA ligase beta subunitInferred from related proteinsTranslation interface
Histidine--tRNA ligaseInferred from related proteinsTranslation interface
Isoleucine--tRNA ligaseInferred from related proteinsTranslation interface
Leucine--tRNA ligase subunit alphaSeen as a proteinTranslation interface
Leucine--tRNA ligase subunit betaInferred from related proteinsTranslation interface
Lysine--tRNA ligaseInferred from related proteinsTranslation interface
Methionine--tRNA ligaseSeen as a proteinTranslation interface
Methionyl-tRNA synthetase beta subunitSeen as a proteinTranslation interface
Phenylalanine--tRNA ligase alpha subunitInferred from related proteinsTranslation interface
Phenylalanine--tRNA ligase beta subunitInferred from related proteinsTranslation interface
Proline--tRNA ligaseSeen as a proteinTranslation interface
Serine--tRNA ligaseSeen as a proteinTranslation interface
Threonine--tRNA ligaseInferred from related proteinsTranslation interface
Tryptophan--tRNA ligaseInferred from related proteinsTranslation interface
Tyrosine--tRNA ligaseInferred from related proteinsTranslation interface
Valine--tRNA ligaseInferred from related proteinsTranslation interface

Epigenetic information

In bacteria, the main epigenetic marks are methyl groups added to DNA, often by restriction–modification systems. The Aquifex genome encodes 2 such DNA methylation or restriction enzymes, and none of them is among the kept proteins. In this scenario, then, the cell carries no DNA-methylation layer. It does keep enzymes that add methyl groups to its RNAs.

Aquifex proteinEvidence (UniProt)Home pathway
Uncharacterized RNA methyltransferase aq_257Inferred from related proteinsNot yet placed in a pathway
tRNA (adenine(58)-N(1))-methyltransferase TrmISeen as a proteinRNA maturation
Probable dual-specificity RNA methyltransferase RlmNInferred from related proteinsNot yet placed in a pathway
Ribosomal RNA small subunit methyltransferase ASeen as a proteinRNA maturation
Ribosomal RNA small subunit methyltransferase ESeen as a proteinRNA maturation
Ribosomal RNA small subunit methyltransferase GInferred from related proteinsRNA maturation
Ribosomal RNA small subunit methyltransferase HInferred from related proteinsRNA maturation
Ribosomal RNA small subunit methyltransferase IInferred from related proteinsRNA maturation
tRNA (guanine(26)-N(2)/guanine(27)-N(2))-dimethyltransferaseSeen as a proteinRNA maturation
tRNA (guanine-N(7)-)-methyltransferaseSeen as a proteinRNA maturation
tRNA (guanine-N(1)-)-methyltransferaseSeen as a proteinRNA maturation
Methylenetetrahydrofolate--tRNA-(uracil-5-)-methyltransferase TrmFOInferred from related proteinsRNA maturation
tRNA (guanosine(18)-2'-O)-methyltransferaseSeen as a proteinRNA maturation

Information not written in the genes

Some of what a cell needs is inherited as structure rather than as sequence. A membrane grows only from an existing membrane, and a ribosome is made with the help of existing ribosomes. A daughter cell therefore starts with working copies of these parts from its parent.

Source: Cavalier-Smith, Trends in Plant Science 5:174–182 (2000)

The origins question

Which came first: the genetic code, or the machinery that reads it?

Naturalistic proposal

Most proposals begin with an RNA stage, in which one kind of molecule both stored information and catalysed reactions. The genetic code is thought to have started with fewer amino acids and codons and to have grown through chemical affinities, selection against costly errors, and later additions.

Experiments have produced ribozymes that copy short RNAs and that attach amino acids to RNA. A system that reads its own code and builds its own readers has not been shown.

Design analysis

The design analysis treats the code as a central case: an arbitrary but consistent mapping between codons and amino acids, enforced by the 22 synthetase proteins kept in the working set.

Every part of the reading machinery is itself written in the code it reads, so the system has to be working in order to produce itself. The design analysis counts this circular dependence as positive evidence for planning. A stepwise, experimentally supported route from chemistry to a coded system would weaken that case.

Hallmarks that apply.

Sources