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Principles of Plant Genetics and Breeding

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52 CHAPTER 3<br />

Enhancer<br />

region<br />

subunits). All genes do not code for proteins, <strong>and</strong> further,<br />

all genes in a cell are not actively transcribing<br />

mRNA all <strong>of</strong> the time. Also, most enzymes are proteins,<br />

but all proteins are not enzymes.<br />

Protein structure<br />

CCAAT box TATA box<br />

Figure 3.19 Diagrammatic presentation <strong>of</strong> a typical eukaryotic gene showing the three basic regions – the upstream 5′<br />

flanking regions, the transcriptional unit, <strong>and</strong> the downstream 3′ flanking region – <strong>and</strong> their constitution.<br />

Polypeptides are precursors <strong>of</strong> proteins. Once produced,<br />

they fold to assume 3D forms, the functional stage that<br />

becomes proteins. There are four basic levels <strong>of</strong> protein<br />

structure – primary, secondary, tertiary, <strong>and</strong> quaternary.<br />

The primary structure <strong>of</strong> proteins is the sequence<br />

<strong>of</strong> the amino acids in the linear backbone <strong>of</strong> the<br />

polypeptide. The next fold (exemplified by the DNA<br />

molecule), is an α-helix, a spiral chain <strong>of</strong> amino acids<br />

stabilized by hydrogen bonds. The secondary structure<br />

describes the arrangement <strong>of</strong> amino acids within certain<br />

areas <strong>of</strong> the polypeptide chain. The tertiary structure is a<br />

3D conformation <strong>of</strong> the entire chain in space. Proteins<br />

with more than one polypeptide chain may exhibit the<br />

quaternary protein structure through aggregations <strong>of</strong><br />

the polypeptides.<br />

Regulation <strong>of</strong> gene expression<br />

Gene regulation is a critical activity performed by<br />

plants for proper growth <strong>and</strong> development. It is not<br />

important for a gene just to be expressed, but its<br />

expression must be regulated such that it is expressed<br />

at the right time only <strong>and</strong> to the desired extent.<br />

Regulation entails the “turning on” <strong>and</strong> “turning <strong>of</strong>f”<br />

5′ non-coding<br />

sequence<br />

Introns<br />

3′ non-coding<br />

sequence<br />

Promoter region Exons Enhancer<br />

region<br />

Upstream 5′ flanking region Transcriptional unit Downstream<br />

3′ flanking<br />

region<br />

<strong>of</strong> genes. It is through regulation <strong>of</strong> gene expression<br />

that cellular adaptation, variation, differentiation, <strong>and</strong><br />

development occur. Some genes are turned on all the<br />

time (called constitutive expression), while others are<br />

turned on only some <strong>of</strong> the time (called differential<br />

expression).<br />

The underlying principle <strong>of</strong> gene regulation is that<br />

there are regulatory molecules that interact with nucleic<br />

acid sequences to control the rate <strong>of</strong> transcription or<br />

translation. Six potential levels for regulation <strong>of</strong> gene<br />

expression exist in eukaryotes – the regulation <strong>of</strong>: (i)<br />

transcription; (ii) RNA processing; (iii) mRNA transport;<br />

(iv) mRNA stability; (v) translation; <strong>and</strong> (vi) protein<br />

activity. Transcription is temporarily <strong>and</strong> spatially<br />

separated from translation in eukaryotes.<br />

A typical eukaryotic gene is shown in Figure 3.19.<br />

Unlike that <strong>of</strong> a monocistronic gene (lacks introns; has<br />

one transcriptional unit <strong>and</strong> one translational unit) as<br />

occurs in bacteria, eukaryotic genes are polycistronic<br />

(split genes with introns). Genes that encode the primary<br />

structures <strong>of</strong> proteins required by all cells for enzymatic<br />

or structural functions are called structural<br />

genes. In prokaryotes, these genes are organized into<br />

clusters that are transcribed as a single unit (coordinately<br />

controlled). The mRNA is called polycistronic mRNA,<br />

coding for multiple proteins involved in the same regulatory<br />

pathway (e.g., the lac operon).<br />

There are two basic categories <strong>of</strong> gene regulation –<br />

negative <strong>and</strong> positive (Figure 3.20). In negative regulation,<br />

an inhibitor that is bound to a DNA (gene) must<br />

be removed in order for transcription to occur. In<br />

positive regulation, gene transcription occurs when an<br />

activator binds to the DNA. One <strong>of</strong> the main ways in

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