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GENE EXPRESSION Gene expression is the process by which information from a gene is used in the synthesis of a functional gene

product. These products are often proteins, but in non-protein coding genes such as ribosomal RNA, transfer RNA or Small nuclear RNA genes, the product is a functional RNA. The process of gene expression is used by all known living organisms to generate the macromolecular machinery for life. GENE EXPRESSION IN EUKARYOTES

Figure 1 showing the summary of gene expression (Wikipedia) Gene expression is virtually the same in all eukaryotes but differences occur with prokaryotes. Plants and animals share basic common mechanisms for example some transcription factors, like

myb and myc factors are similar in structure and function in both plants and animals. Plants and animals contain introns separating the coding regions of most genes and again, they utilize similar machinery to process the introns and form mature mRNAs. Translation in all eukaryotes is also basically the same, though they are some differences relating to process. These mechanisms relate to mRNA structure, including sequences in the 5leader region and those in the 3untranslated region which influence the efficiency and selectivity of translation. TRANSCRIPTION IN EUKARYOTES Transcription is the synthesis of RNA from a DNA template and is catalyzed by enzymes known as RNA polymerase. This process occurs in the nucleus of the eukaryotic cells. During transcription, a DNA sequence is read by RNA polymerase, which produces a complementary, antiparallel RNA strand. RNA polymerase opens up the double strand so that the DNA bases can be accessed to direct the order of bases on the RNA. There are 3 types of RNA polymerase; RNA polymerase 1 responsible for synthesis of rRNA RNA polymerase 11 responsible for synthesis of mRNA RNA polymerase 111 responsible for synthesis of tRNA The eukaryotic RNA polymerase is larger than that of Prokaryotes but their subunits have similar roles. The process of transcription has 3 recognizable steps; initiation, elongation and termination. Initiation This involves recognition of the start point of transcription usually located upstream of the gene of interest. This is determined by a special sequence known as a promoter sequence which is recognized by the RNA polymerase. A Promoter sequence is a unidirectional sequence found on one strand of the DNA that instructs the RNA polymerase in both where to start synthesis and in which direction synthesis should continue. In eukaryotes it is found at -30, -75 and -90 base pairs upstream from the start site of transcription. In addition to promoters, eukaryotic genes also have regulatory regions called enhancers. Both elements (promoter and enhancer) are required for correct expression of eukaryotic genes. As a result of this added complexity, eukaryotic RNA polymerases do not have anything equivalent to the sigma subunit

found in prokaryotic RNA polymerases. Instead, eukaryotes have groups of transcription factors, which are proteins, independent of the RNA polymerases that recognize promoter and enhancer sequences. The most common type of core promoter in eukaryotes is a short DNA sequence known as a TATA box, found -30 base pairs from the start site of transcription. The TATA box is the binding site for a transcription factor known as TATA binding protein (TBP), which is itself a subunit of another transcription factor called Transcription Factor II D (TFIID). TFIID and the other factors (TFIIA, TFIIB, TFIIE, TFIIF, TFIIH, and TFIIJ) form a complex on the DNA that recruits RNA polymerase II to the promoter, and promotes initiation of transcription. These transcription factors are sufficient to get a basal (minimal) level of transcription. Other transcription factors binding to other promoter and enhancer elements are necessary for higher levels of transcription. The completed assembly of transcription factors and RNA polymerase bind to the promoter, forming a transcription initiation complex known as an Open Promoter Complex. Enlongation One strand of DNA, the template strand (or noncoding strand), is used as a template for RNA synthesis. Most RNA molecules start with a G or A and unlike DNA synthesis a primer is not required. As transcription proceeds, RNA polymerase traverses the template strand opening the helix and uses base pairing complimentarity with the DNA template to create an RNA copying in the 3- 5 direction. Only a small scale of DNA is unwound at a time to protect DNA from undesired cleavage. Unlike DNA replication, mRNA transcription can involve multiple RNA polymerases on a single DNA template and multiple rounds of transcription (amplification of particular mRNA), so many mRNA molecules can be rapidly produced from a single copy of a gene. Elongation also involves a proofreading mechanism that can replace incorrectly incorporated bases. In eukaryotes, this may correspond with short pauses during transcription that allow appropriate RNA editing factors to bind. These pauses may be intrinsic to the RNA polymerase or due to chromatin structure. Termination Eukaryotic genes have no strong termination sequences like prokaryotes. Instead, RNA polymerase II continues transcribing up to 1000 to 2000 nucleotides beyond where the 3' end of the mature mRNA will be. The actual 3' end will be determined during RNA processing.

GENE PROCESSING OR EDITING Eukaryotic class II transcripts are processed in order to produce the final mRNA. Processing of the initial transcript includes capping, polyadenylation and intron removal. Ribosomal and transfer RNAs are also processed, but differently; they are neither capped nor poly adenylated. Capping of the RNA

This occurs at the 5' end and a methylated guanine nucleotide is added to the transcript in a 5' to 5' phosphodiester linkage (it's like a nucleotide added to the 5' end in the backwards direction). This 'cap' is important for recognition of the mRNA by ribosomes during translation. Polyadenylation

This involves cleavage of the RNA to produce the proper 3' end, and addition of a string of adenine nucleotides. The position of the 3' end is determined by a sequence within the RNA itself. This sequence, AAUAAA, is known as the polyadenylation signal. When this signal is recognized by the appropriate enzymes, the RNA is cleaved 10 to 30 nucleotides downstream of the signal, and a series of adenine nucleotides is added. Polyadenylation is done without a template, the As are simply added one after another to the 3' end of the RNA. This poly (A) tail, which averages about 200 nucleotides in length, helps protect the RNA from degradation, and plays other regulatory roles. Intron removal

Introns in some RNAs (particularly mitochondrial RNAs) are capable of self-splicing (or autocatalytic splicing). In these splicing reactions, no protein enzyme is required - the enzyme activity resides within the intron RNA itself! Such RNA enzymes are termed ribozymes. Class II RNAs (pre-mRNAs) from most eukaryotes, however, do require protein enzymes to remove their introns. The splicing of these RNAs is carried out by large protein/RNA complex called spliceosomes. Spliceosomes are made up of five different snRNPs called U1, U2, U4, U5, and U6. Each snRNP consists of a specific small nuclear RNA (snRNA) molecule complexed

with protein. Spliceosomes are able to detect intron/exon boundaries, cleave the RNA at the appropriate point, and join adjacent exons together to produce the mature mRNA. Differences between transcription in plants and animals Vertebrate genes can be quite large, often spanning tens of thousands of base pairs and usually separated by numerous large introns, whereas plant genes tend to be much smaller (between 12 kilobases) with fewer and smaller introns. Second, plant transcripts retain introns more often than do animal transcripts (30% of all genes in the model plant, Arabidopsis, compared to 10% in humans). Differences between transcription in Prokaryotes and Eukaryotes In prokaryotes, transcription occurs in the cytoplasm since they dont have a well defined nucleus while in eukaryotes it occurs in the cell's nucleus. Prokaryotic DNA is much more accessible to RNA polymerase than DNA in eukaryotes. Eukaryotic DNA is wrapped around proteins called histones to form structures called nucleosomes. While RNA polymerase interacts directly with prokaryotic DNA, other proteins mediate the interaction between RNA polymerase and DNA in eukaryotes. Prokaryotic RNA polymerase is of one type and made up of 5 polypeptide subunit; 2, and 1 are the catalytic subunits while is the regulatory subunit. It is known as a core enzyme if the sigma unit is missing and a holo enzyme if the sigma unit is present. The sigma subunit is the one that recognizes the promoter sequences -35 and -10 upstream of the gene of interest. Promoter specific initiation in eukaryotes requires more than a dozen basal initiation factors while in prokaryotes it is only a single polypeptide; the sigma subunit. Eukaryotic mRNA contain no ShineDalgarno sequence to show the ribosomes where to start translating instead most eukaryotic mRNA have caps at their 5` which directs initiation factors to bind and begin searching for an initiation codon. Eukaryotic protein synthesis initiation begins with methionine not N formyl methionine Termination of transcription involves formation of a hair pin loop structure and can be dependent or independent of the rho protein factor. Rho-independent terminators have a characteristic structure, which features; a strong G-C rich stem, a hair pin loop and a sequence of 46 U residues in the RNA which are

transcribed from a corresponding stretch of Adenines in the template. Rho-factor-dependent terminators are less well defined. The mRNA produced as a result of transcription is not modified in prokaryotic cells.

TRANSLATION This is the conversion of the information of mRNA into proteins; messenger RNA produced by transcription is decoded by the ribosome to produce a specific amino acid chain, or polypeptide, that will later fold into an active protein. Occurs in the ribosomes which exist as dissociate subunit in the cytoplasm and thus have to be assembled together before translation can take place. The ribosome facilitates decoding by inducing the binding of tRNAs with complementary anticodon sequences to that of the mRNA. The tRNAs carry specific amino acids that are chained together into a polypeptide as the mRNA passes through. It involves several steps; Activation In activation, the correct amino acid is covalently bonded to the correct transfer RNA (tRNA). The amino acid is joined by its carboxyl group to the 3' OH of the tRNA by a peptide bond. When the tRNA has an amino acid linked to it, it is termed "charged". Initiation Initiation of translation usually involves the interaction of certain key proteins with a special tag bound to the 5' cap. The protein factors bind the small ribosomal subunit (40S), and these initiation factors hold the mRNA in place. The eukaryotic Initiation Factor 3 (eIF3) is associated with the small ribosomal subunit, and plays a role in keeping the large ribosomal subunit from prematurely binding. eIF3 also interacts with the eIF4F complex which consists of three other initiation factors: eIF4A, eIF4E and eIF4G. eIF4G is a scaffolding protein which directly associates with both eIF3 and the other two components. eIF4E is the cap-binding protein. It is the rate-limiting step of cap-dependent initiation, and is often cleaved from the complex by some viral proteases to limit the cell's ability to translate its own transcripts.

Elongation With the formation of the complex containing fMet-tRNA in the peptidyl site, an aminoacyl tRNA with the complementary anticodon sequence can bind to the mRNA passing through the acceptor site. This binding is aided by elongation factors that are dependent upon the energy from the hydrolysis of GTP. Elongation factors go through a cycle to regenerate GTP after its hydrolysis. Now, with tRNA bearing a chain of amino acids in the ribosome p site and tRNA containing a single amino acid in the ribosome A site, the addition of a link to the chain can be made. This addition occurs through the formation of a peptide bond, the nitrogen-carbon bond that forms between amino acid subunits to form a polypeptide chain. This bond is catalyzed by the enzyme peptidyl transferase. The peptide bond occurs between the carboxyl group on the lowest link in the peptide chain located at the p site and the amine group on the amino acid in the A group. As a result, the peptide chain shifts over to the A site, with the original amino acid on the A site as the lowest link in the chain. The tRNA in the A site becomes peptidyl RNA, and shifts over to the P site. Meanwhile, the ribosome engages in a process called translocation: spurred by elongation factors, the ribosome moves three nucleotides in the 3' prime direction along the mRNA. In other words, the ribosome moves so that a new mRNA codon is accessible in the A site. Termination Translation ends when one of three stop codons, UAA, UAG, or UGA, enters the A site of the ribosome. There are no aminoacyl tRNA molecules that recognize these sequences. Instead, release factors bind to the P site, catalyzing the release of the completed polypeptide chain and separating the ribosome into its original small and large subunits. POSTTRANSLATIONAL MODIFICATION This is a process which involves chemical modification of a protein after its translation. It is one of the later steps in protein biosynthesis and thus gene expression. During protein synthesis, 20 different amino acids can be incorporated to become a protein. After translation, the posttranslational modification of amino acids extends the range of functions of the protein by attaching it to other biochemical functional groups such as acetate, phosphate, various lipids and carbohydrates, by changing the chemical nature of an amino acid (e.g. citrullination) or by making structural changes, like the formation of disulfide bridges.

Also, enzymes may remove amino acids from the amino end of the protein, or cut the peptide chain in the middle. For instance, the peptide hormone insulin is cut twice after disulfide bonds are formed, and a propeptide is removed from the middle of the chain; the resulting protein consists of two polypeptide chains connected by disulfide bond.

Figure 2 showing the post translation modification in insulin protein (wikipedia) Differences between translation in Eukaryotes and Prokaryotes RNA is usually translated while it is being transcribed since both transcription and translation takes place in the cytoplasm. Transcription of the eukaryotic mRNA occurs in the nucleus, mRNA then moves to the cytoplasm for translation. Another difference is that eukaryotic mRNA must be modified by capping and intron splicing before it can be used to make protein. However, in prokaryotes, the mRNA usually doesnt need much modification and that is why it is able to undergo translation before transcription is done. Most prokaryotic mRNA molecules often contain transcripts of several genes because of clustering of genes related by functions (operons) thus more than one protein is usually expressed by these mRNas. The Eukaryotic mRNA usually specifies only a single protein.

GENETIC CODE The genetic code consists of 64 triplets of nucleotides called codons. A codon is a sequence of 3 bases which specify an amino acid, a start or stop signal.

Characteristics of the genetic code include; Code word

Each codon is a sequence of 3 bases required to determine the incorporation of a single amino acid Colinearity

The order of bases is read in the 3 to 5 direction Degeneracy

Some amino acids have alternate codes or several codons may code for the same amino acids Start codon

It is usually AUG in all species but it also codes for methionine. The codon for methionine is a start signal however in prokaryotes the first methionine is methylated. Stop signal

This is degenerated as it has 3 codons specifying the start signal; UAA, UAG, UGA. This determines the end of the protein once the start codon has been determined, its always the first stop codon encountered down stream. The region between the first methionine and the stop codon is the open reading frame. Universality

This code is the same in all species. In rare incidences this codon is not universal like in animal mitochondria use AUA for methionine not isoleucine, all vertebrate mitochondria use AGA and AGG as chain terminator and in animal and micro organisms mitochondrial UGA codes for Tryptophan while in plants it specifies stop. CODON BIAS All but two of the amino acids (Met and Trp) can be encoded by from 2 to 6 different codons. However, the genome of most organisms reveals that certain codons are preferred over others. In humans, for example, alanine is encoded by GCC four times as often as by GCG. This probably reflects great translation efficiency by the translation apparatus (e.g., ribosomes) for certain codons over their synonyms. REGULATION OF GENE EXPRESSION This refers to the control of the amount and timing of appearance of the functional product of a gene. This is vital to allow a cell to produce the gene products it needs when it needs them; in turn this gives cells the flexibility to adapt to a variable environment, external signals, damage to the cell, etc. More generally gene regulation gives the cell control over all structure and function, and is the basis for cellular differentiation, morphogenesis and the versatility and adaptability of any organism. Any step of gene expression may be modulated, from the DNA-RNA transcription step to posttranslational modification of a protein. The stability of the final gene product contributes to the

expression level of the gene as an unstable product results in a low expression level. Terms used to describe types of genes depending on how they are regulated, these include: A constitutive or house keeping gene is a gene that is transcribed continually. Examples include actin. A facultative gene is a gene which is only transcribed when needed. An inducible gene is a gene whose expression is either responsive to environmental change or dependent on the position in the cell cycle REGULATION OF GENE EXPRESSION IN EUKARYOTES Regulation of gene expression in eukaryotes is complex and occurs on many levels, often overlapping each other. In a hierarchical sense, the chromatin structure it self is the first level influencing gene expression through methylation of select bases, posttranslational modification of histones and alterations in scaffolding. The second level includes; transcription and posttranscriptional processing, including export and turnover from RNAs and the third level of regulation involves translation and posttranslational events.

Pre-transcription regulation In eukaryotes, the accessibility of large regions of DNA can depend on their chromatin structures, which are by histone modifications that are directed by DNA methylation, ncRNA, or DNA-binding protein.

Methylation of DNA by methyltransferase enzymes on cytosine nucleotides in a CpG dinucleotide sequence is a common method of gene silencing. Methylated cytosine residues are unchanged by the treatment, whereas unmethylated ones are changed to uracil. Histone acetylation by Histone acetyl transferase enzymes (HATs) such as CREB-binding protein can also dissociate the DNA from the histone complex thus allowing transcription to proceed. Often, DNA methylation and histone deacetylation work together in gene silencing. The combination of the two seems to be a signal for DNA to be packed more densely, lowering gene expression. Transcription of DNA can be dictated by its structure since the density of its packing can be indicative of the frequency of transcription. Nucleosomes are responsible for the amount of supercoiling of DNA and can be temporarily modified by processes such as phosphorylation or more permanently modified by processes such as methylation. These modifications are considered to be responsible for more or less permanent changes in gene expression levels.

Regulation of transcription This is aimed at controlling how much messenger RNA is transcribed. This stage of gene expression involves activator-enhancer complex which is unique in eukaryotes because they normally have to be activated transcription and includes use of transcription factors and RNA polymerase. Types of transcription factors used include; Activators which enhance the interaction between RNA polymerase and a particular promoter, encouraging the expression of the gene. They increase the attraction of RNA polymerase for the promoter, through interactions with subunits of the RNA polymerase or indirectly by changing the structure of the DNA. Enhancers which are sites on the DNA helix that are bound to by activators in order to loop the DNA bringing a specific promoter to the initiation complex. Enhancers are much more common in eukaryote than prokaryotes. Specificity factors that alter the specificity of RNA polymerase for a given promoter or set of promoters, making it more or less likely to bind to them like the sigma factors used in prokaryotic transcription.

Repressors (silencers) which bind to non-coding sequences on the DNA strand that are close to or overlapping the promoter region, impeding RNA polymerase's progress along the strand, thus impeding the expression of the gene. This is a characteristic of regulation of operons in prokaryotes.

General transcription factors that position RNA polymerase at the start of a proteincoding sequence and then release the polymerase to transcribe the mRNA.

Post transcription regulation After transcription, the RNA must be processed before it can be translated and this presents another level of regulation. RNA processing involves addition of a 5' cap, addition of a 3' poly (A) tail, and removal of introns. Addition of extra nucleotides as a protective cap and tail to the RNA identifies the RNA as an mRNA by the ribosomes, and prevents degradation by cell enzymes as it moves from the nucleus into the cytoplasm. Splicing occurs when introns are removed from the code-carrying nucleotides which are then connected for conversion to proteins. The number of introns regulates the speed at which the RNA can be processed. Regulation can be by either the RNA getting processed or introns being retained in the mRNA. This can determine whether or not an mRNA gets translated. If the RNA is not processed, it will not be transported out of the nucleus and thus will not be translated. The other form of regulation is exon shuffling where some genes have exons that can be exchanged thus affecting the function of the protein produced. An example, a gene with four exons might be spliced differently in two different cell types. In cell 1, exons 1, 2, and 4 would be used in the mRNA:

In cell 2 on the other hand, exons 1, 3, and 4 would be used:

In each of these cases, the polypeptide produced could have a different function. In mammals, for example, the calcitonin gene produces a hormone in one cell type, and a neurotransmitter in another cell type, due to exon shuffling. In Drosophila, alternate splicing of the sex-lethal RNA can produce an mRNA encoding a functional polypeptide, or one with a premature stop codon that encodes a short, nonfunctional polypeptide. Translation Control This can be through controlling the number of ribosomes allowed to attach a single mRNA or controlling the rate at which each ribosome transcribes a message. Another mechanism is use of inhibitory proteins which will prevent the translation of mRNA. They are made inactive when bonded with the substance for which they are trying to block production Regulation of RNA Longevity The longevity of the individual mRNA molecule determines how many times it can be used and reused to create proteins. In eukaryotes, the mRNA tends to be stable thus can be used multiple times; which is efficient, but it prevents eukaryotes from making rapid response changes to environmental disruptions. The mRNA of prokaryotes is unstable, allowing for the creation of new mRNA, which has more opportunities to adjust to environmental conditions. The principle behind regulation of RNA longevity is the difference in the life span of mRNA. Messenger RNAs from different genes have their approximate lifespan encoded in them; this serves to regulate how much of each polypeptide is produced. The information for lifespan is found in the 3' UTR. The sequence AUUUA, when found in the 3' UTR, is a signal for early degradation (and therefore short lifetime). The more times the sequence is present, the shorter the lifespan of the mRNA. Because it is encoded in the nucleotide sequence, this is a set property of each different mRNA; the longevity of an mRNA can't be varied. Post translation control

This will involve the selective cutting and breakdown of proteins that prevent the formation of the final product. In both cases, the hormone or enzyme required to finish or activate the final product may be rendered inactive. CONTROL OF GENE EXPRESSION IN PROKARYOTES In bacteria transcription often occur as polycistrons i.e., many functional-related genes are clustered and transcribed under the same types of regulation known as operons. An operon usually contains regulatory genes and structure genes. Operons can be inducible or repressible. Inducible products are made when the substrate is in the environment (lactose) and needs to be metabolized. Repressible products are made when a signal molecule is scarce like tryptophan. They allow coordinated control of genes required for metabolism. One switch controls more than one gene and they are not present in Eukaryotes. Binding of repressor to operator prevents transcription; RNA polymerase will not be able to bind to promoter. Operators are regions of DNA (15 nucleotides long) that control RNA access to promoter and without repressors transcription will take place. Repressors are regulatory proteins that bind to operator and turn genes off, acting as a braking mechanism (negative control). They are produced at a site away from the operon by a specific regulatory gene. Repressors alternate between active/inactive forms to control transcription. Types of regulatory proteins include; Active form- bound to operator Inactive form- conformation will not allow binding. THE LACTOSE OPERON This is an example of an inducible operon (catabolic). The lac operon is required for the transport and metabolism of lactose in some prokaryotes like E.coli. The cell can use lactose as an energy source by producing the enzyme -galactosidase to digest that lactose into glucose and galactose. However, it would be inefficient to produce enzymes when there is no lactose available or glucose is available. The lac operon uses a two-part control mechanism to ensure that the cell expends energy producing -galactosidase, -galactoside permease and thiogalactoside transacetylase only when necessary. It achieves this with the lac repressor, which halts production in the absence of

lactose, and the Catabolite activator protein (CAP), which assists in production in the absence of glucose. The lac operon consists of three structural genes, and a promoter, a terminator, regulator, and an operator. The three structural genes are: lacZ, lacY, and lacA. lacZ encodes -galactosidase (LacZ), an intracellular enzyme that cleaves the disaccharide lactose into glucose and galactose. lacY encodes -galactoside permease (LacY), a membrane-bound transport protein that pumps lactose into the cell. lacA encodes -galactoside transacetylase (LacA), an enzyme that transfers an acetyl group from acetyl-CoA to -galactosides.

The first control mechanism is the regulatory response to lactose, which uses an intracellular regulatory protein called the lactose repressor to hinder production of -galactosidase in the absence of lactose. The lacI gene coding for the repressor is always expressed (constitutive). If lactose is absent, the repressor binds very tightly to a short DNA sequence just downstream of the promoter near the beginning of lacZ called the lac operator. The repressor binding to the operator interferes with binding of RNA polymerase to the promoter, and therefore mRNA encoding LacZ and LacY is only made at very low levels. In presence of lactose, however, a lactose metabolite called allolactose, binds to the repressor, causing a change in its shape. Thus

altered, the repressor is unable to bind to the operator, allowing RNA polymerase to transcribe the lac genes and thereby leading to high levels of the encoded proteins. The second control mechanism is a response to glucose, which uses the Catabolite activator protein (CAP) to greatly increase production of -galactosidase in the absence of glucose. Cyclic adenosine monophosphate (cAMP) is a signal molecule whose prevalence is inversely proportional to that of glucose. It binds to the CAP, which in turn allows the CAP to bind to the CAP binding site which assists the RNA polymerase in binding to the DNA. In the absence of glucose, the cAMP concentration is high and binding of CAP-cAMP to the DNA significantly increases the production of -galactosidase, enabling the cell to hydrolyse (digest) lactose and release galactose and glucose. SIGNAL TRANSLATION The purposes of signal translation between cells include; regulation of cell growth and differentiation, coordination of different physiological processes and maintainace of homeostasis in stances where circumstances in the environment become stressful. The forms of translation used include; use of molecular messengers or the non-diffusible cell adhesion proteins. MOLECULAR MESSENGERS There are 2 types of molecular messengers that are used to translate signals which affect transcription and include; Hormones Paracrines.

Hormones These are molecules which are released by specialized cells (endocrine glands) into blood and lymph circulation to act on specific cells which may be far away or widely distributed in the body. There are 2 main categories of hormones; steroids and water soluble hormones. a) Steriods

Are small molecular derivatives of cholesterol and grouped into six depending on their physiological action; Androgens, Estrogens, Progestins, Glucocorticoids and mineralocorticoids and Vitamin D

Steroids being lipids in nature are insoluble in blood thus require a carrier proteins for circulation. Since the plasma membrane t is a lipid bilayer, they can freely cross it and behind to specific receptors in the cytoplasm or nucleus (intracellular receptors). The steroid hormones act on DNA directly and may require metabolic transformation before becoming active. The activated hormone- receptor complex then binds to specific regions on the DNA to regulate transcription of various genes. b) Water soluble hormones

They can be peptides or amino acid like hormones. There are hydrophilic thus cannot cross the plasma membrane but need no carrier to move in circulation. They interact with surface receptors only leading to release of a secondary messenger to trigger response. They work by activating intracellular secondary pathways involving cyclic aadenosine monophosphate (cAMP), phospoinositides or calcium-calmodulin in reactions which involves phosporylation of other proteins. Paracrines These are molecular messengers whose action is largely local and restricted to neighboring cells released in small quantities and quickly used up or destroyed. All cell types secrete paracrines, also commonly called growth factors. There are several categories which include; mitogens, trophic factors, chemoattractants and pleiotrophic factors. There are peptides in nature thus some act like the water soluble hormones and bind on the extra cellular receptors. Like hormones they also select their targets on basis of specific receptors. Other growth factors are not freely diffusible thus bind to the extracellular matrix for activity. NON-DIFFUSIBLE ADHESION MOLECULES These are organelles on the cell surface in contact with other cells and extracellular matrix Cell connections involve multiple ligands and cell adhesion receptors. Cell adhesion receptors enable cells to recognize and bind molecules on other cells or in the extracellular matrix. Ligands can be other cell adhesion receptors (receptor-receptor contact) or extracellular matrix molecules. Cell adhesion receptors activate intracellular signaling pathways which will affect transcription. SIGNAL TRANSLATION IN PLANTS

Plants have evolved different signaling mechanisms since plant hormones do not exactly function as do those in animals and partly because plants must cope with environmental changes differently than animals as they cannot physically escape their environments except possibly through reproduction. Therefore, plants have evolved very complex, interacting signaling pathways in response to developmental signals and biotic/abiotic stresses. All of these observations ultimately reflect some of the differences plants display in regulating gene expression compared to yeasts, insects and vertebrates.

REFERENCES "Eukaryotic Chromosomes and Gene Expression." In Biochemistry, 4th ed. New York: W. H. www.emunix.emich.edu/rwinning/genetics/eureg2.htm Stryer, Lubert. "Control of Gene Expression in Prokaryotes." In Biochemistry, 4th ed. New York: W. H. Freeman and Company, 1995 Translation, Paul Billiet St Edwards University Dept and Biochemistry Molecular Biology of the Cell. 4th edition. Alberts B, Johnson A, Lewis J, et al.New York: Garland Science; 2002. Regulation of the mammalian epigenome by long noncoding RNAs by Joanne Whitehead, Gaurav Kumar Pandey and Chandrasekhar. Biochimica et Biophysica Acta (BBA) - General Subjects Volume 1790, Issue 9, September 2009, Pages 936-947 Freeman and Company, 1995