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131-613: Dernburg received her Bachelor of Arts in Biochemistry in 1987 from the University of California, Berkeley . There, she spent half a year working in an organic chemistry lab before she joined Dan Koshland 's laboratory, studying bacterial chemotaxis , or how cells and organisms move in response to a chemical stimulus. Following graduation, she spent a year working as a research technician in Koshland's lab, where she co-authored

262-433: A DNA repair process, and that when it occurs during meiosis it is an adaptation for repairing the genomic DNA that is passed on to progeny. Experimental findings indicate that a substantial benefit of meiosis is recombinational repair of DNA damage in the germline , as indicated by the following examples. Hydrogen peroxide is an agent that causes oxidative stress leading to oxidative DNA damage. Treatment of

393-628: A biological polymer , they undergo a process called dehydration synthesis . Different macromolecules can assemble in larger complexes, often needed for biological activity . Two of the main functions of carbohydrates are energy storage and providing structure. One of the common sugars known as glucose is a carbohydrate, but not all carbohydrates are sugars. There are more carbohydrates on Earth than any other known type of biomolecule; they are used to store energy and genetic information , as well as play important roles in cell to cell interactions and communications . The simplest type of carbohydrate

524-433: A carboxylic acid group, –COOH (although these exist as –NH 3 and –COO under physiologic conditions), a simple hydrogen atom , and a side chain commonly denoted as "–R". The side chain "R" is different for each amino acid of which there are 20 standard ones . It is this "R" group that makes each amino acid different, and the properties of the side chains greatly influence the overall three-dimensional conformation of

655-493: A barrel shaped spindle. In human oocytes spindle microtubule nucleation begins on the chromosomes, forming an aster that eventually expands to surround the chromosomes. Chromosomes then slide along the microtubules towards the equator of the spindle, at which point the chromosome kinetochores form end-on attachments to microtubules. Homologous pairs move together along the metaphase plate: As kinetochore microtubules from both spindle poles attach to their respective kinetochores,

786-399: A chemical theory of metabolism, or even earlier to the 18th century studies on fermentation and respiration by Antoine Lavoisier . Many other pioneers in the field who helped to uncover the layers of complexity of biochemistry have been proclaimed founders of modern biochemistry. Emil Fischer , who studied the chemistry of proteins , and F. Gowland Hopkins , who studied enzymes and

917-465: A common intestinal parasite, was previously considered to have descended from a lineage that predated the emergence of meiosis and sex. However, G. intestinalis has now been found to possess a core set of meiotic genes, including five meiosis specific genes. Also evidence for meiotic recombination , indicative of sexual reproduction , was found in G. intestinalis . Another example of organisms previously thought to be asexual are parasitic protozoa of

1048-434: A component of DNA . A monosaccharide can switch between acyclic (open-chain) form and a cyclic form. The open-chain form can be turned into a ring of carbon atoms bridged by an oxygen atom created from the carbonyl group of one end and the hydroxyl group of another. The cyclic molecule has a hemiacetal or hemiketal group, depending on whether the linear form was an aldose or a ketose . In these cyclic forms,

1179-406: A cyclic [ring] and planar [flat] structure) while others are not. Some are flexible, while others are rigid. Lipids are usually made from one molecule of glycerol combined with other molecules. In triglycerides , the main group of bulk lipids, there is one molecule of glycerol and three fatty acids . Fatty acids are considered the monomer in that case, and maybe saturated (no double bonds in

1310-433: A diploid cell called the zygote . The organism's diploid germ-line stem cells undergo meiosis to make haploid gametes (the spermatozoa in males and ova in females), which fertilize to form the zygote. The diploid zygote undergoes repeated cellular division by mitosis to grow into the organism. In the haplontic life cycle (with post-zygotic meiosis), the organism is haploid, by the proliferation and differentiation of

1441-406: A diploid cell, which contains two copies of each chromosome, termed homologs . First, the cell undergoes DNA replication , so each homolog now consists of two identical sister chromatids. Then each set of homologs pair with each other and exchange genetic information by homologous recombination often leading to physical connections ( crossovers ) between the homologs. In the first meiotic division,

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1572-468: A few (around three to six) monosaccharides are joined, it is called an oligosaccharide ( oligo- meaning "few"). These molecules tend to be used as markers and signals , as well as having some other uses. Many monosaccharides joined form a polysaccharide . They can be joined in one long linear chain, or they may be branched . Two of the most common polysaccharides are cellulose and glycogen , both consisting of repeating glucose monomers . Cellulose

1703-404: A molecule with a 5-membered ring, called glucofuranose . The same reaction can take place between carbons 1 and 5 to form a molecule with a 6-membered ring, called glucopyranose . Cyclic forms with a 7-atom ring called heptoses are rare. Two monosaccharides can be joined by a glycosidic or ester bond into a disaccharide through a dehydration reaction during which a molecule of water

1834-407: A new combination of maternal and paternal genetic information, resulting in offspring that are genetically distinct from either parent. Furthermore, an individual gamete can include an assortment of maternal, paternal, and recombinant chromatids. This genetic diversity resulting from sexual reproduction contributes to the variation in traits upon which natural selection can act. Meiosis uses many of

1965-400: A new diploid organism. The haplodiplontic life cycle can be considered a fusion of the diplontic and haplontic life cycles. Meiosis occurs in all animals and plants. The result, the production of gametes with half the number of chromosomes as the parent cell, is the same, but the detailed process is different. In animals, meiosis produces gametes directly. In land plants and some algae, there

2096-402: A pair of sister chromatids) to opposite poles. Nonkinetochore microtubules lengthen, pushing the centrosomes farther apart. The cell elongates in preparation for division down the center. Unlike in mitosis, only the cohesin from the chromosome arms is degraded while the cohesin surrounding the centromere remains protected by a protein named Shugoshin (Japanese for "guardian spirit"), what prevents

2227-420: A period of rest known as interkinesis or interphase II. No DNA replication occurs during this stage. Meiosis II is the second meiotic division, and usually involves equational segregation, or separation of sister chromatids. Mechanically, the process is similar to mitosis, though its genetic results are fundamentally different. The result is the production of four haploid cells (n chromosomes; 23 in humans) from

2358-423: A prolonged G 2 -like stage known as meiotic prophase . During this time, homologous chromosomes pair with each other and undergo genetic recombination , a programmed process in which DNA may be cut and then repaired, which allows them to exchange some of their genetic information . A subset of recombination events results in crossovers , which create physical links known as chiasmata (singular: chiasma, for

2489-414: A protein. Some amino acids have functions by themselves or in a modified form; for instance, glutamate functions as an important neurotransmitter . Amino acids can be joined via a peptide bond . In this dehydration synthesis, a water molecule is removed and the peptide bond connects the nitrogen of one amino acid's amino group to the carbon of the other's carboxylic acid group. The resulting molecule

2620-566: A reducing end because of full acetal formation between the aldehyde carbon of glucose (C1) and the keto carbon of fructose (C2). Lipids comprise a diverse range of molecules and to some extent is a catchall for relatively water-insoluble or nonpolar compounds of biological origin, including waxes , fatty acids , fatty-acid derived phospholipids , sphingolipids , glycolipids , and terpenoids (e.g., retinoids and steroids ). Some lipids are linear, open-chain aliphatic molecules, while others have ring structures. Some are aromatic (with

2751-399: A single haploid cell called the gamete . Two organisms of opposing sex contribute their haploid gametes to form a diploid zygote. The zygote undergoes meiosis immediately, creating four haploid cells. These cells undergo mitosis to create the organism. Many fungi and many protozoa utilize the haplontic life cycle. In the haplodiplontic life cycle (with sporic or intermediate meiosis),

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2882-626: A study analyzing the structure of a bacterial sensory receptor. Dernburg then entered the Tetrad Program at the University of California, San Francisco for her doctoral work. She received her PhD in 1996 working in the laboratory of John Sedat studying several aspects of chromosomes organization and function. She developed fluorescence in situ hybridization (FISH) methods to study the genetic content of cells and to investigate how chromosomes are organized within cell nuclei. Using these tools she investigated how chromosome organization within

3013-440: A study of the components and composition of living things and how they come together to become life. In this sense, the history of biochemistry may therefore go back as far as the ancient Greeks . However, biochemistry as a specific scientific discipline began sometime in the 19th century, or a little earlier, depending on which aspect of biochemistry is being focused on. Some argued that the beginning of biochemistry may have been

3144-448: A total of four daughter cells, each with a haploid set of chromosomes. Meiosis is now complete and ends up with four new daughter cells. Meiosis appears to be a fundamental characteristic of eukaryotic organisms and to have been present early in eukaryotic evolution. Eukaryotes that were once thought to lack meiotic sex have recently been shown to likely have, or once have had, this capability. As one example, Giardia intestinalis ,

3275-479: Is crossed over , creating new combinations of code on each chromosome. Later on, during fertilisation , the haploid cells produced by meiosis from a male and a female will fuse to create a zygote , a cell with two copies of each chromosome again. Errors in meiosis resulting in aneuploidy (an abnormal number of chromosomes) are the leading known cause of miscarriage and the most frequent genetic cause of developmental disabilities . In meiosis, DNA replication

3406-431: Is a monosaccharide , which among other properties contains carbon , hydrogen , and oxygen , mostly in a ratio of 1:2:1 (generalized formula C n H 2 n O n , where n is at least 3). Glucose (C 6 H 12 O 6 ) is one of the most important carbohydrates; others include fructose (C 6 H 12 O 6 ), the sugar commonly associated with the sweet taste of fruits , and deoxyribose (C 5 H 10 O 4 ),

3537-416: Is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes , the sperm or egg cells . It involves two rounds of division that ultimately result in four cells, each with only one copy of each chromosome ( haploid ). Additionally, prior to the division, genetic material from the paternal and maternal copies of each chromosome

3668-400: Is an alternation of generations such that meiosis in the diploid sporophyte generation produces haploid spores instead of gametes. When they germinate, these spores undergo repeated cell division by mitosis, developing into a multicellular haploid gametophyte generation, which then produces gametes directly (i.e. without further meiosis). In both animals and plants, the final stage is for

3799-412: Is an energy source in most life forms. For instance, polysaccharides are broken down into their monomers by enzymes ( glycogen phosphorylase removes glucose residues from glycogen, a polysaccharide). Disaccharides like lactose or sucrose are cleaved into their two component monosaccharides. Glucose is mainly metabolized by a very important ten-step pathway called glycolysis , the net result of which

3930-407: Is an important structural component of plant's cell walls and glycogen is used as a form of energy storage in animals. Sugar can be characterized by having reducing or non-reducing ends. A reducing end of a carbohydrate is a carbon atom that can be in equilibrium with the open-chain aldehyde ( aldose ) or keto form ( ketose ). If the joining of monomers takes place at such a carbon atom,

4061-424: Is called a dipeptide , and short stretches of amino acids (usually, fewer than thirty) are called peptides or polypeptides . Longer stretches merit the title proteins . As an example, the important blood serum protein albumin contains 585 amino acid residues . Proteins can have structural and/or functional roles. For instance, movements of the proteins actin and myosin ultimately are responsible for

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4192-423: Is cleaved, allowing the sister chromatids to segregate. The sister chromatids by convention are now called sister chromosomes as they move toward opposing poles. The process ends with telophase II , which is similar to telophase I, and is marked by decondensation and lengthening of the chromosomes and the disassembly of the spindle. Nuclear envelopes re-form and cleavage or cell plate formation eventually produces

4323-520: Is divided into meiosis I and meiosis II which are further divided into Karyokinesis I, Cytokinesis I, Karyokinesis II, and Cytokinesis II, respectively. The preparatory steps that lead up to meiosis are identical in pattern and name to interphase of the mitotic cell cycle. Interphase is divided into three phases: Interphase is followed by meiosis I and then meiosis II. Meiosis I separates replicated homologous chromosomes, each still made up of two sister chromatids, into two daughter cells, thus reducing

4454-624: Is followed by two rounds of cell division to produce four daughter cells, each with half the number of chromosomes as the original parent cell. The two meiotic divisions are known as meiosis I and meiosis II . Before meiosis begins, during S phase of the cell cycle , the DNA of each chromosome is replicated so that it consists of two identical sister chromatids , which remain held together through sister chromatid cohesion. This S-phase can be referred to as "premeiotic S-phase" or "meiotic S-phase". Immediately following DNA replication, meiotic cells enter

4585-578: Is likely mediated by oxidative stress leading to increased DNA damage. Meiosis occurs in eukaryotic life cycles involving sexual reproduction , consisting of the cyclical process of growth and development by mitotic cell division, production of gametes by meiosis and fertilization. At certain stages of the life cycle, germ cells produce gametes. Somatic cells make up the body of the organism and are not involved in gamete production. Cycling meiosis and fertilization events results in alternation between haploid and diploid states. The organism phase of

4716-421: Is not an essential element for plants. Plants need boron and silicon , but animals may not (or may need ultra-small amounts). Just six elements— carbon , hydrogen , nitrogen , oxygen , calcium and phosphorus —make up almost 99% of the mass of living cells, including those in the human body (see composition of the human body for a complete list). In addition to the six major elements that compose most of

4847-508: Is released. The reverse reaction in which the glycosidic bond of a disaccharide is broken into two monosaccharides is termed hydrolysis . The best-known disaccharide is sucrose or ordinary sugar , which consists of a glucose molecule and a fructose molecule joined. Another important disaccharide is lactose found in milk, consisting of a glucose molecule and a galactose molecule. Lactose may be hydrolysed by lactase , and deficiency in this enzyme results in lactose intolerance . When

4978-506: Is the generic name of the family of biopolymers . They are complex, high-molecular-weight biochemical macromolecules that can convey genetic information in all living cells and viruses. The monomers are called nucleotides , and each consists of three components: a nitrogenous heterocyclic base (either a purine or a pyrimidine ), a pentose sugar, and a phosphate group. The most common nucleic acids are deoxyribonucleic acid (DNA) and ribonucleic acid (RNA). The phosphate group and

5109-449: Is the stage at which all autosomal chromosomes have synapsed. In this stage homologous recombination, including chromosomal crossover (crossing over), is completed through the repair of the double strand breaks formed in leptotene. Most breaks are repaired without forming crossovers resulting in gene conversion . However, a subset of breaks (at least one per chromosome) form crossovers between non-sister (homologous) chromosomes resulting in

5240-537: Is the study of chemical processes within and relating to living organisms . A sub-discipline of both chemistry and biology , biochemistry may be divided into three fields: structural biology , enzymology , and metabolism . Over the last decades of the 20th century, biochemistry has become successful at explaining living processes through these three disciplines. Almost all areas of the life sciences are being uncovered and developed through biochemical methodology and research. Biochemistry focuses on understanding

5371-492: Is thought to be the reason why complex life appeared only after Earth's atmosphere accumulated large amounts of oxygen. In vertebrates , vigorously contracting skeletal muscles (during weightlifting or sprinting, for example) do not receive enough oxygen to meet the energy demand, and so they shift to anaerobic metabolism , converting glucose to lactate. The combination of glucose from noncarbohydrates origin, such as fat and proteins. This only happens when glycogen supplies in

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5502-409: Is to break down one molecule of glucose into two molecules of pyruvate . This also produces a net two molecules of ATP , the energy currency of cells, along with two reducing equivalents of converting NAD (nicotinamide adenine dinucleotide: oxidized form) to NADH (nicotinamide adenine dinucleotide: reduced form). This does not require oxygen; if no oxygen is available (or the cell cannot use oxygen),

5633-412: The University of California, Berkeley to continue investigating chromosome organization and dynamics, focusing on meiosis using the nematode worm Caenorhabditis elegans as a model organism . Her laboratory has contributed to the community's understanding of how chromosomes find and pair with the appropriate homolog during meiosis , which is essential for proper chromosome segregation and ensuring

5764-423: The amino acids , which are used to synthesize proteins ). The mechanisms used by cells to harness energy from their environment via chemical reactions are known as metabolism . The findings of biochemistry are applied primarily in medicine , nutrition and agriculture . In medicine, biochemists investigate the causes and cures of diseases . Nutrition studies how to maintain health and wellness and also

5895-521: The chemistry required for biological activity of molecules, molecular biology studies their biological activity, genetics studies their heredity, which happens to be carried by their genome . This is shown in the following schematic that depicts one possible view of the relationships between the fields: Meiosis This is an accepted version of this page Meiosis ( / m aɪ ˈ oʊ s ɪ s / ; from Ancient Greek μείωσις ( meíōsis )  'lessening', (since it

6026-440: The diakinesis stage, from Greek words meaning "moving through". This is the first point in meiosis where the four parts of the tetrads are actually visible. Sites of crossing over entangle together, effectively overlapping, making chiasmata clearly visible. Other than this observation, the rest of the stage closely resembles prometaphase of mitosis; the nucleoli disappear, the nuclear membrane disintegrates into vesicles, and

6157-416: The essential amino acids . Mammals do possess the enzymes to synthesize alanine , asparagine , aspartate , cysteine , glutamate , glutamine , glycine , proline , serine , and tyrosine , the nonessential amino acids. While they can synthesize arginine and histidine , they cannot produce it in sufficient amounts for young, growing animals, and so these are often considered essential amino acids. If

6288-424: The glutamate residue at position 6 with a valine residue changes the behavior of hemoglobin so much that it results in sickle-cell disease . Finally, quaternary structure is concerned with the structure of a protein with multiple peptide subunits, like hemoglobin with its four subunits. Not all proteins have more than one subunit. Ingested proteins are usually broken up into single amino acids or dipeptides in

6419-761: The him-8 gene, which encodes a protein responsible for proper meiotic separation of the worm's X chromosome. These proteins facilitate an interaction between the pairing centers and a complex of microtubules and a motor protein called dynein , which allow chromosomes to move along the nuclear envelope until they encounter their partner. Dernburg has also moved beyond the nematode worm to begin to study meiotic mechanisms in planarians , which are non-parasitic flatworms, as well as another species of nematode called Pristionchus pacificus to understand how meiotic mechanisms are conserved—or diverge—across species. The Dernburg lab also develops methods for microscopy of living nematodes. Using live cell imaging they discovered that

6550-478: The meiotic spindle begins to form. Unlike mitotic cells, human and mouse oocytes do not have centrosomes to produce the meiotic spindle. In mice, approximately 80 MicroTubule Organizing Centers (MTOCs) form a sphere in the ooplasm and begin to nucleate microtubules that reach out towards chromosomes, attaching to the chromosomes at the kinetochore . Over time, the MTOCs merge until two poles have formed, generating

6681-549: The small intestine and then absorbed. They can then be joined to form new proteins. Intermediate products of glycolysis, the citric acid cycle, and the pentose phosphate pathway can be used to form all twenty amino acids, and most bacteria and plants possess all the necessary enzymes to synthesize them. Humans and other mammals, however, can synthesize only half of them. They cannot synthesize isoleucine , leucine , lysine , methionine , phenylalanine , threonine , tryptophan , and valine . Because they must be ingested, these are

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6812-530: The synaptonemal complex , the special structure that holds together homologous chromosomes and enables them to undergo meiotic recombination , is a liquid crystal . This has led them to propose a mechanism for crossover interference based on a reaction–diffusion mechanism. In 2008, Dernburg became a Howard Hughes Medical Institute Investigator. She also has a joint appointment as a Faculty Senior Scientist at Lawrence Berkeley National Laboratory . Biochemistry Biochemistry or biological chemistry

6943-399: The " vital principle ") distinct from any found in non-living matter, and it was thought that only living beings could produce the molecules of life. In 1828, Friedrich Wöhler published a paper on his serendipitous urea synthesis from potassium cyanate and ammonium sulfate ; some regarded that as a direct overthrow of vitalism and the establishment of organic chemistry . However,

7074-449: The DNA of the germline . The repair process used appears to involve homologous recombinational repair Prophase I arrested oocytes have a high capability for efficient repair of DNA damage , particularly exogenously induced double-strand breaks. DNA repair capability appears to be a key quality control mechanism in the female germ line and a critical determinant of fertility . Genetic recombination can be viewed as fundamentally

7205-554: The DNA are required for pairing and synapsis between homologous chromosomes during meiosis. Dernburg found that in Caenorhabditis elegans , double-strand breaks are required for recombination and for chromosome segregation during meiosis, but not for homologous pairing and synapsis. The finding suggested that there may be more diversity in meiotic mechanisms than was previously expected. In 2000, Dernburg started up her laboratory at Lawrence Berkeley National Laboratory and

7336-502: The Greek letter Chi , Χ) between the homologous chromosomes. In most organisms, these links can help direct each pair of homologous chromosomes to segregate away from each other during meiosis I, resulting in two haploid cells that have half the number of chromosomes as the parent cell. During meiosis II, the cohesion between sister chromatids is released and they segregate from one another, as during mitosis . In some cases, all four of

7467-518: The N-terminal domain. The enzyme-linked immunosorbent assay (ELISA), which uses antibodies, is one of the most sensitive tests modern medicine uses to detect various biomolecules. Probably the most important proteins, however, are the enzymes . Virtually every reaction in a living cell requires an enzyme to lower the activation energy of the reaction. These molecules recognize specific reactant molecules called substrates ; they then catalyze

7598-492: The NAD is restored by converting the pyruvate to lactate (lactic acid) (e.g. in humans) or to ethanol plus carbon dioxide (e.g. in yeast ). Other monosaccharides like galactose and fructose can be converted into intermediates of the glycolytic pathway. In aerobic cells with sufficient oxygen , as in most human cells, the pyruvate is further metabolized. It is irreversibly converted to acetyl-CoA , giving off one carbon atom as

7729-485: The Wöhler synthesis has sparked controversy as some reject the death of vitalism at his hands. Since then, biochemistry has advanced, especially since the mid-20th century, with the development of new techniques such as chromatography , X-ray diffraction , dual polarisation interferometry , NMR spectroscopy , radioisotopic labeling , electron microscopy and molecular dynamics simulations. These techniques allowed for

7860-409: The ability to reproduce by parthenogenesis . Meiosis does not occur in archaea or bacteria , which generally reproduce asexually via binary fission . However, a "sexual" process known as horizontal gene transfer involves the transfer of DNA from one bacterium or archaeon to another and recombination of these DNA molecules of different parental origin. Meiosis was discovered and described for

7991-416: The amino group is removed from an amino acid, it leaves behind a carbon skeleton called an α- keto acid . Enzymes called transaminases can easily transfer the amino group from one amino acid (making it an α-keto acid) to another α-keto acid (making it an amino acid). This is important in the biosynthesis of amino acids, as for many of the pathways, intermediates from other biochemical pathways are converted to

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8122-434: The amount of energy gained from glycolysis (six molecules of ATP are used, compared to the two gained in glycolysis). Analogous to the above reactions, the glucose produced can then undergo glycolysis in tissues that need energy, be stored as glycogen (or starch in plants), or be converted to other monosaccharides or joined into di- or oligosaccharides. The combined pathways of glycolysis during exercise, lactate's crossing via

8253-878: The animals' needs. Unicellular organisms release the ammonia into the environment. Likewise, bony fish can release ammonia into the water where it is quickly diluted. In general, mammals convert ammonia into urea, via the urea cycle . In order to determine whether two proteins are related, or in other words to decide whether they are homologous or not, scientists use sequence-comparison methods. Methods like sequence alignments and structural alignments are powerful tools that help scientists identify homologies between related molecules. The relevance of finding homologies among proteins goes beyond forming an evolutionary pattern of protein families . By finding how similar two protein sequences are, we acquire knowledge about their structure and therefore their function. Nucleic acids , so-called because of their prevalence in cellular nuclei ,

8384-436: The appropriate chromosome copy number in daughter cells . Her group has worked to understand how special regions of chromosomes, known as pairing centers, promote homologous chromosome pairing, synapsis, and segregation in the worm. In 2005, they published a study demonstrating how pairing centers perform two separable functions during meiosis. First, they facilitate pairing through stabilizing an intermediate complex involved in

8515-468: The bloodstream to the liver, subsequent gluconeogenesis and release of glucose into the bloodstream is called the Cori cycle . Researchers in biochemistry use specific techniques native to biochemistry, but increasingly combine these with techniques and ideas developed in the fields of genetics , molecular biology , and biophysics . There is not a defined line between these disciplines. Biochemistry studies

8646-513: The body and are broken into fatty acids and glycerol, the final degradation products of fats and lipids. Lipids, especially phospholipids , are also used in various pharmaceutical products , either as co-solubilizers (e.g. in parenteral infusions) or else as drug carrier components (e.g. in a liposome or transfersome ). Proteins are very large molecules—macro-biopolymers—made from monomers called amino acids . An amino acid consists of an alpha carbon atom attached to an amino group, –NH 2 ,

8777-539: The carbon chain) or unsaturated (one or more double bonds in the carbon chain). Most lipids have some polar character and are largely nonpolar. In general, the bulk of their structure is nonpolar or hydrophobic ("water-fearing"), meaning that it does not interact well with polar solvents like water . Another part of their structure is polar or hydrophilic ("water-loving") and will tend to associate with polar solvents like water. This makes them amphiphilic molecules (having both hydrophobic and hydrophilic portions). In

8908-490: The case of cholesterol , the polar group is a mere –OH (hydroxyl or alcohol). In the case of phospholipids, the polar groups are considerably larger and more polar, as described below. Lipids are an integral part of our daily diet. Most oils and milk products that we use for cooking and eating like butter , cheese , ghee etc. are composed of fats . Vegetable oils are rich in various polyunsaturated fatty acids (PUFA). Lipid-containing foods undergo digestion within

9039-410: The chemical basis which allows biological molecules to give rise to the processes that occur within living cells and between cells, in turn relating greatly to the understanding of tissues and organs as well as organism structure and function. Biochemistry is closely related to molecular biology , the study of the molecular mechanisms of biological phenomena. Much of biochemistry deals with

9170-535: The chromosome number by half. During meiosis II, sister chromatids decouple and the resultant daughter chromosomes are segregated into four daughter cells. For diploid organisms, the daughter cells resulting from meiosis are haploid and contain only one copy of each chromosome. In some species, cells enter a resting phase known as interkinesis between meiosis I and meiosis II. Meiosis I and II are each divided into prophase , metaphase , anaphase , and telophase stages, similar in purpose to their analogous subphases in

9301-596: The chromosomes at the first meiotic division. The paired and replicated chromosomes are called bivalents (two chromosomes) or tetrads (four chromatids ), with one chromosome coming from each parent. Prophase I is divided into a series of substages which are named according to the appearance of chromosomes. The first stage of prophase I is the leptotene stage, also known as leptonema , from Greek words meaning "thin threads". In this stage of prophase I, individual chromosomes—each consisting of two replicated sister chromatids—become "individualized" to form visible strands within

9432-507: The contraction of skeletal muscle. One property many proteins have is that they specifically bind to a certain molecule or class of molecules—they may be extremely selective in what they bind. Antibodies are an example of proteins that attach to one specific type of molecule. Antibodies are composed of heavy and light chains. Two heavy chains would be linked to two light chains through disulfide linkages between their amino acids. Antibodies are specific through variation based on differences in

9563-591: The discovery and detailed analysis of many molecules and metabolic pathways of the cell , such as glycolysis and the Krebs cycle (citric acid cycle), and led to an understanding of biochemistry on a molecular level. Another significant historic event in biochemistry is the discovery of the gene , and its role in the transfer of information in the cell. In the 1950s, James D. Watson , Francis Crick , Rosalind Franklin and Maurice Wilkins were instrumental in solving DNA structure and suggesting its relationship with

9694-492: The discovery of the first enzyme , diastase (now called amylase ), in 1833 by Anselme Payen , while others considered Eduard Buchner 's first demonstration of a complex biochemical process alcoholic fermentation in cell-free extracts in 1897 to be the birth of biochemistry. Some might also point as its beginning to the influential 1842 work by Justus von Liebig , Animal chemistry, or, Organic chemistry in its applications to physiology and pathology , which presented

9825-519: The dynamic nature of biochemistry, represent two examples of early biochemists. The term "biochemistry" was first used when Vinzenz Kletzinsky (1826–1882) had his "Compendium der Biochemie" printed in Vienna in 1858; it derived from a combination of biology and chemistry . In 1877, Felix Hoppe-Seyler used the term ( biochemie in German) as a synonym for physiological chemistry in the foreword to

9956-429: The effect of chromosomal positioning on gene expression. Dernburg also studied the role heterochromatin plays in chromosome segregation during meiosis, finding that the heterochromatic regions of homologous chromosomes remain associated with one another until metaphase I , or the stage at which chromosomes line up along the center of the nucleus prior to the first round of meiotic division. This association ensures that

10087-549: The effects of nutritional deficiencies . In agriculture, biochemists investigate soil and fertilizers with the goal of improving crop cultivation, crop storage, and pest control . In recent decades, biochemical principles and methods have been combined with problem-solving approaches from engineering to manipulate living systems in order to produce useful tools for research, industrial processes, and diagnosis and control of disease—the discipline of biotechnology . At its most comprehensive definition, biochemistry can be seen as

10218-554: The electrons from high-energy states in NADH and quinol is conserved first as proton gradient and converted to ATP via ATP synthase. This generates an additional 28 molecules of ATP (24 from the 8 NADH + 4 from the 2 quinols), totaling to 32 molecules of ATP conserved per degraded glucose (two from glycolysis + two from the citrate cycle). It is clear that using oxygen to completely oxidize glucose provides an organism with far more energy than any oxygen-independent metabolic feature, and this

10349-426: The enzyme can be regulated, enabling control of the biochemistry of the cell as a whole. The structure of proteins is traditionally described in a hierarchy of four levels. The primary structure of a protein consists of its linear sequence of amino acids; for instance, "alanine-glycine-tryptophan-serine-glutamate-asparagine-glycine-lysine-...". Secondary structure is concerned with local morphology (morphology being

10480-469: The enzyme complexes of the respiratory chain, an electron transport system transferring the electrons ultimately to oxygen and conserving the released energy in the form of a proton gradient over a membrane ( inner mitochondrial membrane in eukaryotes). Thus, oxygen is reduced to water and the original electron acceptors NAD and quinone are regenerated. This is why humans breathe in oxygen and breathe out carbon dioxide. The energy released from transferring

10611-469: The exchange of genetic information. The exchange of information between the homologous chromatids results in a recombination of information; each chromosome has the complete set of information it had before, and there are no gaps formed as a result of the process. Because the chromosomes cannot be distinguished in the synaptonemal complex, the actual act of crossing over is not perceivable through an ordinary light microscope, and chiasmata are not visible until

10742-448: The first issue of Zeitschrift für Physiologische Chemie (Journal of Physiological Chemistry) where he argued for the setting up of institutes dedicated to this field of study. The German chemist Carl Neuberg however is often cited to have coined the word in 1903, while some credited it to Franz Hofmeister . It was once generally believed that life and its materials had some essential property or substance (often referred to as

10873-632: The first time in sea urchin eggs in 1876 by the German biologist Oscar Hertwig . It was described again in 1883, at the level of chromosomes , by the Belgian zoologist Edouard Van Beneden , in Ascaris roundworm eggs. The significance of meiosis for reproduction and inheritance, however, was described only in 1890 by German biologist August Weismann , who noted that two cell divisions were necessary to transform one diploid cell into four haploid cells if

11004-451: The flies their red eyes; when the brown gene is turned off, flies have brown eyes. Heterochromatic regions of chromosomes tend to associate together in a specific compartment of the nucleus. Dernburg found that when a region of heterochromatin was inserted near the brown gene, the gene would associate with a heterochromatic region of the nucleus and would thus be inactivated, giving flies with this insertion brown eyes. The study demonstrated

11135-477: The formation of sex cells with the wrong number of chromosomes. Dernburg's group also discovered that the functions of pairing centers depend on a family of four DNA-binding zinc-finger proteins —called HIM or ZIM proteins—that recognize and bind short, repetitive sequences that are hallmarks of pairing centers. Each him or zim protein recognizes a particular pairing center sequence to help bring homologous chromosomes together. Dernburg's group first uncovered

11266-467: The free hydroxy group of the pyranose or furanose form is exchanged with an OH-side-chain of another sugar, yielding a full acetal . This prevents opening of the chain to the aldehyde or keto form and renders the modified residue non-reducing. Lactose contains a reducing end at its glucose moiety, whereas the galactose moiety forms a full acetal with the C4-OH group of glucose. Saccharose does not have

11397-482: The genetic material of the cell, nucleic acids often play a role as second messengers , as well as forming the base molecule for adenosine triphosphate (ATP), the primary energy-carrier molecule found in all living organisms. Also, the nitrogenous bases possible in the two nucleic acids are different: adenine, cytosine, and guanine occur in both RNA and DNA, while thymine occurs only in DNA and uracil occurs in RNA. Glucose

11528-526: The genetic transfer of information. In 1958, George Beadle and Edward Tatum received the Nobel Prize for work in fungi showing that one gene produces one enzyme . In 1988, Colin Pitchfork was the first person convicted of murder with DNA evidence, which led to the growth of forensic science . More recently, Andrew Z. Fire and Craig C. Mello received the 2006 Nobel Prize for discovering

11659-443: The genus Leishmania , which cause human disease. However, these organisms were shown to have a sexual cycle consistent with a meiotic process. Although amoeba were once generally regarded as asexual, evidence has been presented that most lineages are anciently sexual and that the majority of asexual groups probably arose recently and independently. Dacks and Rogers proposed, based on a phylogenetic analysis, that facultative sex

11790-518: The homologous chromosomes become much more closely (~100 nm) and stably paired (a process called synapsis) mediated by the installation of the transverse and central elements of the synaptonemal complex . Synapsis is thought to occur in a zipper-like fashion starting from a recombination nodule. The paired chromosomes are called bivalent or tetrad chromosomes. The pachytene stage ( / ˈ p æ k ɪ t iː n / PAK -i-teen ), also known as pachynema , from Greek words meaning "thick threads".

11921-414: The homologous chromosomes, forming inter-axis bridges, and resulting in the pairing/co-alignment of homologues (to a distance of ~400 nm in mice). Leptotene is followed by the zygotene stage, also known as zygonema , from Greek words meaning "paired threads", which in some organisms is also called the bouquet stage because of the way the telomeres cluster at one end of the nucleus. In this stage

12052-460: The homologs are segregated to separate daughter cells by the spindle apparatus . The cells then proceed to a second division without an intervening round of DNA replication. The sister chromatids are segregated to separate daughter cells to produce a total of four haploid cells. Female animals employ a slight variation on this pattern and produce one large ovum and three small polar bodies. Because of recombination, an individual chromatid can consist of

12183-448: The human body, humans require smaller amounts of possibly 18 more. The 4 main classes of molecules in biochemistry (often called biomolecules ) are carbohydrates , lipids , proteins , and nucleic acids . Many biological molecules are polymers : in this terminology, monomers are relatively small macromolecules that are linked together to create large macromolecules known as polymers. When monomers are linked together to synthesize

12314-402: The idiosyncratic rendering "maiosis": We propose to apply the terms Maiosis or Maiotic phase to cover the whole series of nuclear changes included in the two divisions that were designated as Heterotype and Homotype by Flemming . The spelling was changed to "meiosis" by Koernicke (1905) and by Pantel and De Sinety (1906) to follow the usual conventions for transliterating Greek . Meiosis

12445-491: The laboratory of Anne Villeneuve at Stanford University , where she transitioned to working on the nematode worm Caenorhabditis elegans . There, focused on the process of meiosis, which she continues to study in her own lab, she adapted FISH methods to study the cytology of chromosome pairing in the worm. In 1998, she published a study documenting how meiosis in the worm is distinct from meiosis in many other eukaryotic organisms . In most eukaryotes, double-strand breaks in

12576-417: The life cycle can occur either during the diploid state ( diplontic life cycle), during the haploid state ( haplontic life cycle), or both ( haplodiplontic life cycle), in which there are two distinct organism phases, one with haploid cells and the other with diploid cells. In the diplontic life cycle (with pre-gametic meiosis), as in humans, the organism is multicellular and diploid, grown by mitosis from

12707-458: The liver are worn out. The pathway is a crucial reversal of glycolysis from pyruvate to glucose and can use many sources like amino acids, glycerol and Krebs Cycle . Large scale protein and fat catabolism usually occur when those suffer from starvation or certain endocrine disorders. The liver regenerates the glucose, using a process called gluconeogenesis . This process is not quite the opposite of glycolysis, and actually requires three times

12838-458: The living organism alternates between haploid and diploid states. Consequently, this cycle is also known as the alternation of generations . The diploid organism's germ-line cells undergo meiosis to produce spores. The spores proliferate by mitosis, growing into a haploid organism. The haploid organism's gamete then combines with another haploid organism's gamete, creating the zygote. The zygote undergoes repeated mitosis and differentiation to produce

12969-556: The meiotic products form gametes such as sperm , spores or pollen . In female animals, three of the four meiotic products are typically eliminated by extrusion into polar bodies , and only one cell develops to produce an ovum . Because the number of chromosomes is halved during meiosis, gametes can fuse (i.e. fertilization ) to form a diploid zygote that contains two copies of each chromosome, one from each parent. Thus, alternating cycles of meiosis and fertilization enable sexual reproduction , with successive generations maintaining

13100-577: The metaphase plate during metaphase I and orientation of sister chromatids in metaphase II, this is the subsequent separation of homologs and sister chromatids during anaphase I and II, it allows a random and independent distribution of chromosomes to each daughter cell (and ultimately to gametes); and (2) Crossing Over . The physical exchange of homologous chromosomal regions by homologous recombination during prophase I results in new combinations of genetic information within chromosomes. However, such physical exchange does not always occur during meiosis. In

13231-704: The metaphase plate, with respect to the orientation of the other bivalents along the same equatorial line. The protein complex cohesin holds sister chromatids together from the time of their replication until anaphase. In mitosis, the force of kinetochore microtubules pulling in opposite directions creates tension. The cell senses this tension and does not progress with anaphase until all the chromosomes are properly bi-oriented. In meiosis, establishing tension ordinarily requires at least one crossover per chromosome pair in addition to cohesin between sister chromatids (see Chromosome segregation ). Kinetochore microtubules shorten, pulling homologous chromosomes (which each consist of

13362-431: The mitotic cell cycle. Therefore, meiosis includes the stages of meiosis I (prophase I, metaphase I, anaphase I, telophase I) and meiosis II (prophase II, metaphase II, anaphase II, telophase II). During meiosis, specific genes are more highly transcribed . In addition to strong meiotic stage-specific expression of mRNA , there are also pervasive translational controls (e.g. selective usage of preformed mRNA), regulating

13493-417: The next stage. During the diplotene stage, also known as diplonema , from Greek words meaning "two threads", the synaptonemal complex disassembles and homologous chromosomes separate from one another a little. However, the homologous chromosomes of each bivalent remain tightly bound at chiasmata, the regions where crossing-over occurred. The chiasmata remain on the chromosomes until they are severed at

13624-415: The nucleus can affect transcription , and how chromosomes interact to separate from each during the process of meiosis , using the fly Drosophila as a model organism . Specifically, she used FISH to monitor the chromosomal position of regions of heterochromatin —a tightly packed region of DNA associated with non-expressed genes—and of a fly gene called brown , which codes for the red pigment that gives

13755-452: The nucleus. The chromosomes each form a linear array of loops mediated by cohesin , and the lateral elements of the synaptonemal complex assemble forming an "axial element" from which the loops emanate. Recombination is initiated in this stage by the enzyme SPO11 which creates programmed double strand breaks (around 300 per meiosis in mice). This process generates single stranded DNA filaments coated by RAD51 and DMC1 which invade

13886-479: The number of chromosomes had to be maintained. In 1911, the American geneticist Thomas Hunt Morgan detected crossovers in meiosis in the fruit fly Drosophila melanogaster , which helped to establish that genetic traits are transmitted on chromosomes. The term "meiosis" is derived from the Greek word μείωσις , meaning 'lessening'. It was introduced to biology by J.B. Farmer and J.E.S. Moore in 1905, using

14017-417: The oocytes of the silkworm Bombyx mori , meiosis is completely achiasmate (lacking crossovers). Although synaptonemal complexes are present during the pachytene stage of meiosis in B. mori , crossing-over homologous recombination is absent between the paired chromosomes . Female mammals and birds are born possessing all the oocytes needed for future ovulations, and these oocytes are arrested at

14148-417: The paired homologous chromosomes align along an equatorial plane that bisects the spindle, due to continuous counterbalancing forces exerted on the bivalents by the microtubules emanating from the two kinetochores of homologous chromosomes. This attachment is referred to as a bipolar attachment. The physical basis of the independent assortment of chromosomes is the random orientation of each bivalent along with

14279-466: The pairing process. Second, pairing centers promote the formation of a synaptonemal complex , in which a protein polymer acts as a scaffold to hold homologous chromosomes together during recombination. In a related study, her group also uncovered a conserved meiotic checkpoint that acts during meiosis to recognize unpaired/unsynapsed chromosomes. Cells identified as having unsynapsed chromosomes undergo apoptosis , or programmed cell death, to guard against

14410-399: The pinching of the cell membrane in animal cells or the formation of the cell wall in plant cells, occurs, completing the creation of two daughter cells. However, cytokinesis does not fully complete resulting in "cytoplasmic bridges" which enable the cytoplasm to be shared between daughter cells until the end of meiosis II. Sister chromatids remain attached during telophase I. Cells may enter

14541-730: The ploidy is reduced from diploid to haploid, meiosis I is referred to as a reductional division . Meiosis II is an equational division analogous to mitosis, in which the sister chromatids are segregated, creating four haploid daughter cells (1n, 1c). Prophase I is by far the longest phase of meiosis (lasting 13 out of 14 days in mice ). During prophase I, homologous maternal and paternal chromosomes pair, synapse , and exchange genetic information (by homologous recombination ), forming at least one crossover per chromosome. These crossovers become visible as chiasmata (plural; singular chiasma ). This process facilitates stable pairing between homologous chromosomes and hence enables accurate segregation of

14672-447: The prophase I stage of meiosis. In humans, as an example, oocytes are formed between three and four months of gestation within the fetus and are therefore present at birth. During this prophase I arrested stage ( dictyate ), which may last for decades, four copies of the genome are present in the oocytes. The arrest of ooctyes at the four genome copy stage was proposed to provide the informational redundancy needed to repair damage in

14803-410: The reaction between them. By lowering the activation energy , the enzyme speeds up that reaction by a rate of 10 or more; a reaction that would normally take over 3,000 years to complete spontaneously might take less than a second with an enzyme. The enzyme itself is not used up in the process and is free to catalyze the same reaction with a new set of substrates. Using various modifiers, the activity of

14934-651: The resulting daughter cells have the appropriate number of chromosomes. She completed her dissertation, called Nuclear Architecture in Drosophila melanogaster , documenting this work in 1996. Her thesis received the Larry Sandler Memorial Award in 1997 from the Genetics Society of America , which recognizes the most outstanding dissertation in the area of Drosophila genetics and biology. For her postdoctoral research, Dernburg joined

15065-413: The ring usually has 5 or 6 atoms. These forms are called furanoses and pyranoses , respectively—by analogy with furan and pyran , the simplest compounds with the same carbon-oxygen ring (although they lack the carbon-carbon double bonds of these two molecules). For example, the aldohexose glucose may form a hemiacetal linkage between the hydroxyl on carbon 1 and the oxygen on carbon 4, yielding

15196-559: The role of RNA interference (RNAi) in the silencing of gene expression . Around two dozen chemical elements are essential to various kinds of biological life . Most rare elements on Earth are not needed by life (exceptions being selenium and iodine ), while a few common ones ( aluminum and titanium ) are not used. Most organisms share element needs, but there are a few differences between plants and animals . For example, ocean algae use bromine , but land plants and animals do not seem to need any. All animals require sodium , but

15327-401: The same mechanisms as mitosis , the type of cell division used by eukaryotes to divide one cell into two identical daughter cells. In some plants, fungi, and protists meiosis results in the formation of spores : haploid cells that can divide vegetatively without undergoing fertilization. Some eukaryotes, like bdelloid rotifers , do not have the ability to carry out meiosis and have acquired

15458-561: The same number of chromosomes, occurs in all organisms that utilize meiosis. Meiosis occurs in all sexually-reproducing single-celled and multicellular organisms (which are all eukaryotes ), including animals , plants and fungi . It is an essential process for oogenesis and spermatogenesis . Although the process of meiosis is related to the more general cell division process of mitosis , it differs in two important respects: usually occurs between identical sister chromatids and does not result in genetic changes Meiosis begins with

15589-465: The same number of chromosomes. For example, diploid human cells contain 23 pairs of chromosomes including 1 pair of sex chromosomes (46 total), half of maternal origin and half of paternal origin. Meiosis produces haploid gametes (ova or sperm) that contain one set of 23 chromosomes. When two gametes (an egg and a sperm) fuse, the resulting zygote is once again diploid, with the mother and father each contributing 23 chromosomes. This same pattern, but not

15720-405: The second meiotic division. In metaphase II , the centromeres contain two kinetochores that attach to spindle fibers from the centrosomes at opposite poles. The new equatorial metaphase plate is rotated by 90 degrees when compared to meiosis I, perpendicular to the previous plate. This is followed by anaphase II , in which the remaining centromeric cohesin, not protected by Shugoshin anymore,

15851-452: The sister chromatids from separating. This allows the sister chromatids to remain together while homologs are segregated. The first meiotic division effectively ends when the chromosomes arrive at the poles. Each daughter cell now has half the number of chromosomes but each chromosome consists of a pair of chromatids. The microtubules that make up the spindle network disappear, and a new nuclear membrane surrounds each haploid set. Cytokinesis,

15982-419: The structures, functions, and interactions of biological macromolecules such as proteins , nucleic acids , carbohydrates , and lipids . They provide the structure of cells and perform many of the functions associated with life. The chemistry of the cell also depends upon the reactions of small molecules and ions . These can be inorganic (for example, water and metal ions) or organic (for example,

16113-476: The study of structure). Some combinations of amino acids will tend to curl up in a coil called an α-helix or into a sheet called a β-sheet ; some α-helixes can be seen in the hemoglobin schematic above. Tertiary structure is the entire three-dimensional shape of the protein. This shape is determined by the sequence of amino acids. In fact, a single change can change the entire structure. The alpha chain of hemoglobin contains 146 amino acid residues; substitution of

16244-683: The sugar of each nucleotide bond with each other to form the backbone of the nucleic acid, while the sequence of nitrogenous bases stores the information. The most common nitrogenous bases are adenine , cytosine , guanine , thymine , and uracil . The nitrogenous bases of each strand of a nucleic acid will form hydrogen bonds with certain other nitrogenous bases in a complementary strand of nucleic acid. Adenine binds with thymine and uracil, thymine binds only with adenine, and cytosine and guanine can bind only with one another. Adenine, thymine, and uracil contain two hydrogen bonds, while hydrogen bonds formed between cytosine and guanine are three. Aside from

16375-446: The transition to anaphase I to allow homologous chromosomes to move to opposite poles of the cell. In human fetal oogenesis , all developing oocytes develop to this stage and are arrested in prophase I before birth. This suspended state is referred to as the dictyotene stage or dictyate. It lasts until meiosis is resumed to prepare the oocyte for ovulation, which happens at puberty or even later. Chromosomes condense further during

16506-428: The two haploid cells (with n chromosomes, each consisting of two sister chromatids) produced in meiosis I. The four main steps of meiosis II are: prophase II, metaphase II, anaphase II, and telophase II. In prophase II , we see the disappearance of the nucleoli and the nuclear envelope again as well as the shortening and thickening of the chromatids. Centrosomes move to the polar regions and arrange spindle fibers for

16637-413: The ultimate meiotic stage-specific protein expression of genes during meiosis. Thus, both transcriptional and translational controls determine the broad restructuring of meiotic cells needed to carry out meiosis. Meiosis I segregates homologous chromosomes , which are joined as tetrads (2n, 4c), producing two haploid cells (n chromosomes, 23 in humans) which each contain chromatid pairs (1n, 2c). Because

16768-425: The waste product carbon dioxide , generating another reducing equivalent as NADH . The two molecules acetyl-CoA (from one molecule of glucose) then enter the citric acid cycle , producing two molecules of ATP, six more NADH molecules and two reduced (ubi)quinones (via FADH 2 as enzyme-bound cofactor), and releasing the remaining carbon atoms as carbon dioxide. The produced NADH and quinol molecules then feed into

16899-401: The yeast Schizosaccharomyces pombe with hydrogen peroxide increased the frequency of mating and the formation of meiotic spores by 4 to 18-fold. Volvox carteri , a haploid multicellular, facultatively sexual green algae, can be induced by heat shock to reproduce by meiotic sex. This induction can be inhibited by antioxidants indicating that the induction of meiotic sex by heat shock

17030-474: The α-keto acid skeleton, and then an amino group is added, often via transamination . The amino acids may then be linked together to form a protein. A similar process is used to break down proteins. It is first hydrolyzed into its component amino acids. Free ammonia (NH3), existing as the ammonium ion (NH4+) in blood, is toxic to life forms. A suitable method for excreting it must therefore exist. Different tactics have evolved in different animals, depending on

17161-404: Was likely present in the common ancestor of eukaryotes. The new combinations of DNA created during meiosis are a significant source of genetic variation alongside mutation, resulting in new combinations of alleles , which may be beneficial. Meiosis generates gamete genetic diversity in two ways: (1) Law of Independent Assortment . The independent orientation of homologous chromosome pairs along

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