Category: Curriculum Map

Chapter 12    The Cell Cycle    Lecture Outline

Overview

·         The ability of organisms to reproduce their kind is the one characteristic that best distinguishes living things from nonliving matter.

·         The continuity of life is based on the reproduction of cells, or cell division.

A. The Key Roles of Cell Division

1. Cell division functions in reproduction, growth, and repair.

·         The division of a unicellular organism reproduces an entire organism, increasing the population.

·         Cell division on a larger scale can produce progeny for some multicellular organisms.

°         This includes organisms that can grow by cuttings.

·         Cell division enables a multicellular organism to develop from a single fertilized egg or zygote.

·         In a multicellular organism, cell division functions to repair and renew cells that die from normal wear and tear or accidents.

·         Cell division is part of the cell cycle, the life of a cell from its origin in the division of a parent cell until its own division into two.

2. Cell division results in genetically identical daughter cells.

·         Cell division requires the distribution of identical genetic material—DNA—to two daughter cells.

·         What is remarkable is the fidelity with which DNA is passed along, without dilution, from one generation to the next.

·         A dividing cell duplicates its DNA, allocates the two copies to opposite ends of the cell, and then splits into two daughter cells.

·         A cell’s genetic information, packaged as DNA, is called its genome.

°         In prokaryotes, the genome is often a single long DNA molecule.

°         In eukaryotes, the genome consists of several DNA molecules.

·         A human cell must duplicate about 2 m of DNA and separate the two copies such that each daughter cell ends up with a complete genome.

·         DNA molecules are packaged into chromosomes.

°         Every eukaryotic species has a characteristic number of chromosomes in each cell nucleus.

§            Human somatic cells (body cells) have 46 chromosomes, made up of two sets of 23 (one from each parent).

§            Human gametes (sperm or eggs) have one set of 23 chromosomes, half the number in a somatic cell.

·         Eukaryotic chromosomes are made of chromatin, a complex of DNA and associated protein.

°         Each single chromosome contains one long, linear DNA molecule carrying hundreds or thousands of genes, the units that specify an organism’s inherited traits.

·         The associated proteins maintain the structure of the chromosome and help control gene activity.

·         When a cell is not dividing, each chromosome is in the form of a long, thin chromatin fiber.

·         Before cell division, chromatin condenses, coiling and folding to make a smaller package.

·         Each duplicated chromosome consists of two sister chromatids, which contain identical copies of the chromosome’s DNA.

°         The chromatids are initially attached by adhesive proteins along their lengths.

°         As the chromosomes condense, the region where the chromatids connect shrinks to a narrow area, the centromere.

·         Later in cell division, the sister chromatids are pulled apart and repackaged into two new nuclei at opposite ends of the parent cell.

°         Once the sister chromatids separate, they are considered individual chromosomes.

·         Mitosis, the formation of the two daughter nuclei, is usually followed by division of the cytoplasm, cytokinesis.

·         These processes start with one cell and produce two cells that are genetically identical to the original parent cell.

°         Each of us inherited 23 chromosomes from each parent: one set in an egg and one set in sperm.

°         The fertilized egg, or zygote, underwent cycles of mitosis and cytokinesis to produce a fully developed multicellular human made up of 200 trillion somatic cells.

°         These processes continue every day to replace dead and damaged cells.

°         Essentially, these processes produce clones—cells with identical genetic information.

·         In contrast, gametes (eggs or sperm) are produced only in gonads (ovaries or testes) by a variation of cell division called meiosis.

°         Meiosis yields four nonidentical daughter cells, each with half the chromosomes of the parent.

°         In humans, meiosis reduces the number of chromosomes from 46 to 23.

°         Fertilization fuses two gametes together and doubles the number of chromosomes to 46 again.

B. The Mitotic Cell Cycle

1. The mitotic phase alternates with interphase in the cell cycle.

·         The mitotic (M) phase of the cell cycle alternates with the much longer interphase.

°         The M phase includes mitosis and cytokinesis.

°         Interphase accounts for 90% of the cell cycle.

·         During interphase, the cell grows by producing proteins and cytoplasmic organelles, copies its chromosomes, and prepares for cell division.

·         Interphase has three subphases: the G1 phase (“first gap”), the S phase (“synthesis”), and the G2 phase (“second gap”).

°         During all three subphases, the cell grows by producing proteins and cytoplasmic organelles such as mitochondria and endoplasmic reticulum.

°         However, chromosomes are duplicated only during the S phase.

·         The daughter cells may then repeat the cycle.

·         A typical human cell might divide once every 24 hours.

°         Of this time, the M phase would last less than an hour, while the S phase might take 10–12 hours, or half the cycle.

°         The rest of the time would be divided between the G1 and G2 phases.

°         The G1 phase varies most in length from cell to cell.

·         Mitosis is a continuum of changes.

·         For convenience, mitosis is usually broken into five subphases: prophase, prometaphase, metaphase, anaphase, and telophase.

·         In late interphase, the chromosomes have been duplicated but are not condensed.

°         A nuclear membrane bounds the nucleus, which contains one or more nucleoli.

°         The centrosome has replicated to form two centrosomes.

°         In animal cells, each centrosome features two centrioles.

·         In prophase, the chromosomes are tightly coiled, with sister chromatids joined together.

°         The nucleoli disappear.

°         The mitotic spindle begins to form.

§            It is composed of centrosomes and the microtubules that extend from them.

°         The radial arrays of shorter microtubules that extend from the centrosomes are called asters.

°         The centrosomes move away from each other, apparently propelled by lengthening microtubules.

·         During prometaphase, the nuclear envelope fragments, and microtubules from the spindle interact with the condensed chromosomes.

°         Each of the two chromatids of a chromosome has a kinetochore, a specialized protein structure located at the centromere.

°         Kinetochore microtubules from each pole attach to one of two kinetochores.

°         Nonkinetochore microtubules interact with those from opposite ends of the spindle.

·         The spindle fibers push the sister chromatids until they are all arranged at the metaphase plate, an imaginary plane equidistant from the poles, defining metaphase.

·         At anaphase, the centromeres divide, separating the sister chromatids.

°         Each is now pulled toward the pole to which it is attached by spindle fibers.

°         By the end, the two poles have equivalent collections of chromosomes.

·         At telophase, daughter nuclei begin to form at the two poles.

°         Nuclear envelopes arise from the fragments of the parent cell’s nuclear envelope and other portions of the endomembrane system.

°         The chromosomes become less tightly coiled.

·         Cytokinesis, division of the cytoplasm, is usually well underway by late telophase.

°         In animal cells, cytokinesis involves the formation of a cleavage furrow, which pinches the cell in two.

°         In plant cells, vesicles derived from the Golgi apparatus produce a cell plate at the middle of the cell.

2. The mitotic spindle distributes chromosomes to daughter cells: a closer look.

·         The mitotic spindle, fibers composed of microtubules and associated proteins, is a major driving force in mitosis.

·         As the spindle assembles during prophase, the elements come from partial disassembly of the cytoskeleton.

·         The spindle fibers elongate by incorporating more subunits of the protein tubulin.

·         Assembly of the spindle microtubules starts in the centrosome.

°         The centrosome (microtubule-organizing center) is a nonmembranous organelle that organizes the cell’s microtubules.

°         In animal cells, the centrosome has a pair of centrioles at the center, but the centrioles are not essential for cell division.

·         During interphase, the single centrosome replicates to form two centrosomes.

·         As mitosis starts, the two centrosomes are located near the nucleus.

°         As the spindle microtubules grow from them, the centrioles are pushed apart.

°         By the end of prometaphase, they are at opposite ends of the cell.

·         An aster, a radial array of short microtubules, extends from each centrosome.

·         The spindle includes the centrosomes, the spindle microtubules, and the asters.

·         Each sister chromatid has a kinetochore of proteins and chromosomal DNA at the centromere.

°         The kinetochores of the joined sister chromatids face in opposite directions.

·         During prometaphase, some spindle microtubules (called kinetochore microtubules) attach to the kinetochores.

·         When a chromosome’s kinetochore is “captured” by microtubules, the chromosome moves toward the pole from which those microtubules come.

·         When microtubules attach to the other pole, this movement stops and a tug-of-war ensues.

·         Eventually, the chromosome settles midway between the two poles of the cell, on the metaphase plate.

·         Nonkinetochore microtubules from opposite poles overlap and interact with each other.

·         By metaphase, the microtubules of the asters have grown and are in contact with the plasma membrane.

·         The spindle is now complete.

·         Anaphase commences when the proteins holding the sister chromatids together are inactivated.

°         Once the chromosomes are separate, full-fledged chromosomes, they move toward opposite poles of the cell.

·         How do the kinetochore microtubules function into the poleward movement of chromosomes?

·         One hypothesis is that the chromosomes are “reeled in” by the shortening of microtubules at the spindle poles.

·         Experimental evidence supports the hypothesis that motor proteins on the kinetochore “walk” the attached chromosome along the microtubule toward the nearest pole.

°         Meanwhile, the excess microtubule sections depolymerize at their kinetochore ends.

·         What is the function of the nonkinetochore microtubules?

·         Nonkinetochore microtubules are responsible for lengthening the cell along the axis defined by the poles.

°         These microtubules interdigitate and overlap across the metaphase plate.

°         During anaphase, the area of overlap is reduced as motor proteins attached to the microtubules walk them away from one another, using energy from ATP.

°         As microtubules push apart, the microtubules lengthen by the addition of new tubulin monomers to their overlapping ends, allowing continued overlap.

3. Cytokinesis divides the cytoplasm: a closer look.

·         Cytokinesis, division of the cytoplasm, typically follows mitosis.

·         In animal cells, cytokinesis occurs by a process called cleavage.

·         The first sign of cleavage is the appearance of a cleavage furrow in the cell surface near the old metaphase plate.

·         On the cytoplasmic side of the cleavage furrow is a contractile ring of actin microfilaments associated with molecules of the motor protein myosin.

°         Contraction of the ring pinches the cell in two.

·         Cytokinesis in plants, which have cell walls, involves a completely different mechanism.

·         During telophase, vesicles from the Golgi coalesce at the metaphase plate, forming a cell plate.

°         The plate enlarges until its membranes fuse with the plasma membrane at the perimeter.

°         The contents of the vesicles form new cell wall material between the daughter cells.

4. Mitosis in eukaryotes may have evolved from binary fission in bacteria.

·         Prokaryotes reproduce by binary fission, not mitosis.

·         Most bacterial genes are located on a single bacterial chromosome that consists of a circular DNA molecule and associated proteins.

·         While bacteria are smaller and simpler than eukaryotic cells, they still have large amounts of DNA that must be copied and distributed equally to two daughter cells.

·         The circular bacterial chromosome is highly folded and coiled in the cell.

·         In binary fission, chromosome replication begins at one point in the circular chromosome, the origin of replication site, producing two origins.

°         As the chromosome continues to replicate, one origin moves toward each end of the cell.

°         While the chromosome is replicating, the cell elongates.

°         When replication is complete, its plasma membrane grows inward to divide the parent cell into two daughter cells, each with a complete genome.

·         Researchers have developed methods to allow them to observe the movement of bacterial chromosomes.

°         The movement is similar to the poleward movements of the centromere regions of eukaryotic chromosomes.

°         However, bacterial chromosomes lack visible mitotic spindles or even microtubules.

·         The mechanism behind the movement of the bacterial chromosome is becoming clearer but is still not fully understood.

°         Several proteins have been identified and play important roles.

·         How did mitosis evolve?

°         There is evidence that mitosis had its origins in bacterial binary fission.

°         Some of the proteins involved in binary fission are related to eukaryotic proteins.

°         Two of these are related to eukaryotic tubulin and actin proteins.

·         As eukaryotes evolved, the ancestral process of binary fission gave rise to mitosis.

·         Possible intermediate evolutionary steps are seen in the division of two types of unicellular algae.

°         In dinoflagellates, replicated chromosomes are attached to the nuclear envelope.

°         In diatoms, the spindle develops within the nucleus.

·         In most eukaryotic cells, the nuclear envelope breaks down and a spindle separates the chromosomes.

C. Regulation of the Cell Cycle

·         The timing and rates of cell division in different parts of an animal or plant are crucial for normal growth, development, and maintenance.

·         The frequency of cell division varies with cell type.

°         Some human cells divide frequently throughout life (skin cells).

°         Others have the ability to divide, but keep it in reserve (liver cells).

°         Mature nerve and muscle cells do not appear to divide at all after maturity.

·         Investigation of the molecular mechanisms regulating these differences provide important insights into the operation of normal cells, and may also explain cancer cells escape controls.

1. Cytoplasmic signals drive the cell cycle.

·         The cell cycle appears to be driven by specific chemical signals present in the cytoplasm.

·         Some of the initial evidence for this hypothesis came from experiments in which cultured mammalian cells at different phases of the cell cycle were fused to form a single cell with two nuclei.

°         Fusion of an S phase cell and a G1 phase cell induces the G1 nucleus to start S phase.

§            This suggests that chemicals present in the S phase nucleus stimulated the fused cell.

°         Fusion of a cell in mitosis (M phase) with one in interphase (even G1 phase) induces the second cell to enter mitosis.

·         The sequential events of the cell cycle are directed by a distinct cell cycle control system.

°         Cyclically operating molecules trigger and coordinate key events in the cell cycle.

°         The control cycle has a built-in clock, but it is also regulated by external adjustments and internal controls.

·         A checkpoint in the cell cycle is a critical control point where stop and go-ahead signals regulate the cycle.

°         The signals are transmitted within the cell by signal transduction pathways.

°         Animal cells generally have built-in stop signals that halt the cell cycle at checkpoints until overridden by go-ahead signals.

°         Many signals registered at checkpoints come from cellular surveillance mechanisms.

°         These indicate whether key cellular processes have been completed correctly.

°         Checkpoints also register signals from outside the cell.

·         Three major checkpoints are found in the G1, G2, and M phases.

·         For many cells, the G1 checkpoint, the “restriction point” in mammalian cells, is the most important.

°         If the cell receives a go-ahead signal at the G1 checkpoint, it usually completes the cell cycle and divides.

°         If it does not receive a go-ahead signal, the cell exits the cycle and switches to a nondividing state, the G0 phase.

§            Most cells in the human body are in this phase.

§            Liver cells can be “called back” to the cell cycle by external cues, such as growth factors released during injury.

§            Highly specialized nerve and muscle cells never divide.

·         Rhythmic fluctuations in the abundance and activity of cell cycle control molecules pace the events of the cell cycle.

°         These regulatory molecules include protein kinases that activate or deactivate other proteins by phosphorylating them.

·         These kinases are present in constant amounts but require attachment of a second protein, a cyclin, to become activated.

°         Levels of cyclin proteins fluctuate cyclically.

°         Because of the requirement for binding of a cyclin, the kinases are called cyclin-dependent kinases, or Cdks.

·         Cyclin levels rise sharply throughout interphase, and then fall abruptly during mitosis.

·         Peaks in the activity of one cyclin-Cdk complex, MPF, correspond to peaks in cyclin concentration.

·         MPF (“maturation-promoting factor” or “M-phase-promoting-factor”) triggers the cell’s passage past the G2 checkpoint to the M phase.

°         MPF promotes mitosis by phosphorylating a variety of other protein kinases.

°         MPF stimulates fragmentation of the nuclear envelope by phosphorylation of various proteins of the nuclear lamina.

°         It also triggers the breakdown of cyclin, dropping cyclin and MPF levels during mitosis and inactivating MPF.

§            The noncyclin part of MPF, the Cdk, persists in the cell in inactive form until it associates with new cyclin molecules synthesized during the S and G2 phases of the next round of the cycle.

·         At least three Cdk proteins and several cyclins regulate the key G1 checkpoint.

·         Similar mechanisms are also involved in driving the cell cycle past the M phase checkpoint.

2. Internal and external cues help regulate the cell cycle.

·         While research scientists know that active Cdks function by phosphorylating proteins, the identity of all these proteins is still under investigation.

·         Scientists do not yet know what Cdks actually do in most cases.

·         Some steps in the signaling pathways that regulate the cell cycle are clear.

°         Some signals originate inside the cell, others outside.

·         The M phase checkpoint ensures that all the chromosomes are properly attached to the spindle at the metaphase plate before anaphase.

°         This ensures that daughter cells do not end up with missing or extra chromosomes.

·         A signal to delay anaphase originates at kinetochores that have not yet attached to spindle microtubules.

°         This keeps the anaphase-promoting complex (APC) in an inactive state.

°         When all kinetochores are attached, the APC activates, triggering breakdown of cyclin and inactivation of proteins holding sister chromatids together.

·         A variety of external chemical and physical factors can influence cell division.

°         For example, cells fail to divide if an essential nutrient is left out of the culture medium.

·         Particularly important for mammalian cells are growth factors, proteins released by one group of cells that stimulate other cells to divide.

°         For example, platelet-derived growth factors (PDGF), produced by platelet blood cells, bind to tyrosine-kinase receptors of fibroblasts, a type of connective tissue cell.

°         This triggers a signal-transduction pathway that allows cells to pass the G1 checkpoint and divide.

·         Each cell type probably responds specifically to a certain growth factor or combination of factors.

·         The role of PDGF is easily seen in cell culture.

°         Fibroblasts in culture will only divide in the presence of a medium that also contains PDGF.

·         In a living organism, platelets release PDGF in the vicinity of an injury.

°         The resulting proliferation of fibroblasts helps heal the wound.

·         At least 50 different growth factors can trigger specific cells to divide.

·         The effect of an external physical factor on cell division can be seen in density-dependent inhibition of cell division.

°         Cultured cells normally divide until they form a single layer on the inner surface of the culture container.

°         If a gap is created, the cells will grow to fill the gap.

°         At high densities, the amount of growth factors and nutrients is insufficient to allow continued cell growth.

·         Most animal cells also exhibit anchorage dependence for cell division.

°         To divide, they must be anchored to a substratum, typically the extracellular matrix of a tissue.

°         Control appears to be mediated by pathways involving plasma membrane proteins and elements of the cytoskeleton linked to them.

·         Cancer cells exhibit neither density-dependent inhibition nor anchorage dependence.

3. Cancer cells have escaped from cell cycle controls.

·         Cancer cells divide excessively and invade other tissues because they are free of the body’s control mechanisms.

°         Cancer cells do not stop dividing when growth factors are depleted.

°         This is either because a cancer cell manufactures its own growth factors, has an abnormality in the signaling pathway, or has an abnormal cell cycle control system.

·         If and when cancer cells stop dividing, they do so at random points, not at the normal checkpoints in the cell cycle.

·         Cancer cells may divide indefinitely if they have a continual supply of nutrients.

°         In contrast, nearly all mammalian cells divide 20 to 50 times under culture conditions before they stop, age, and die.

·         Cancer cells may be “immortal.”

°         HeLa cells from a tumor removed from a woman (Henrietta Lacks) in 1951 are still reproducing in culture.

·         The abnormal behavior of cancer cells begins when a single cell in a tissue undergoes a transformation that converts it from a normal cell to a cancer cell.

°         Normally, the immune system recognizes and destroys transformed cells.

°         However, cells that evade destruction proliferate to form a tumor, a mass of abnormal cells.

·         If the abnormal cells remain at the originating site, the lump is called a benign tumor.

°         Most do not cause serious problems and can be fully removed by surgery.

·         In a malignant tumor, the cells become invasive enough to impair the functions of one or more organs.

·         In addition to chromosomal and metabolic abnormalities, cancer cells often lose attachment to nearby cells, are carried by the blood and lymph system to other tissues, and start more tumors in an event called metastasis.

°         Cancer cells are abnormal in many ways.

°         They may have an unusual number of chromosomes, their metabolism may be disabled, and they may cease to function in any constructive way.

°         Cancer cells may secrete signal molecules that cause blood vessels to grow toward the tumor.

·         Treatments for metastasizing cancers include high-energy radiation and chemotherapy with toxic drugs.

°         These treatments target actively dividing cells.

°         Chemotherapeutic drugs interfere with specific steps in the cell cycle.

°         For example, Taxol prevents mitotic depolymerization, preventing cells from proceeding past metaphase.

°         The side effects of chemotherapy are due to the drug’s effects on normal cells.

·         Researchers are beginning to understand how a normal cell is transformed into a cancer cell.

°         The causes are diverse, but cellular transformation always involves the alteration of genes that influence the cell cycle control system.

 

 

Chapter 13    Meiosis and Sexual Life Cycles    Lecture Outline

Overview

·         Living organisms are distinguished by their ability to reproduce their own kind.

·         Offspring resemble their parents more than they do less closely related individuals of the same species.

·         The transmission of traits from one generation to the next is called heredity or inheritance.

·         However, offspring differ somewhat from parents and siblings, demonstrating variation.

·         Farmers have bred plants and animals for desired traits for thousands of years, but the mechanisms of heredity and variation eluded biologists until the development of genetics in the 20th century.

·         Genetics is the scientific study of heredity and variation.

A. The Basis of Heredity

1. Offspring acquire genes from parents by inheriting chromosomes.

·         Parents endow their offspring with coded information in the form of genes.

°         Your genome is comprosed of the tens of thousands of genes that you inherited from your mother and your father.

·         Genes program specific traits that emerge as we develop from fertilized eggs into adults.

·         Genes are segments of DNA. Genetic information is transmitted as specific sequences of the four deoxyribonucleotides in DNA.

°         This is analogous to the symbolic information of language in which words and sentences are translated into mental images.

°         Cells translate genetic “sentences” into freckles and other features with no resemblance to genes.

·         Most genes program cells to synthesize specific enzymes and other proteins whose cumulative action produces an organism’s inherited traits.

·         The transmission of hereditary traits has its molecular basis in the precise replication of DNA.

°         This produces copies of genes that can be passed from parents to offspring.

·         In plants and animals, sperm and ova (unfertilized eggs) transmit genes from one generation to the next.

·         After fertilization (fusion of a sperm cell and an ovum), genes from both parents are present in the nucleus of the fertilized egg, or zygote.

·         Almost all the DNA in a eukaryotic cell is subdivided into chromosomes in the nucleus.

°         Tiny amounts of DNA are also found in mitochondria and chloroplasts.

·         Every living species has a characteristic number of chromosomes.

°         Humans have 46 chromosomes in almost all of their cells.

·         Each chromosome consists of a single DNA molecule associated with various proteins.

·         Each chromosome has hundreds or thousands of genes, each at a specific location, its locus.

2. Like begets like, more or less: a comparison of asexual and sexual reproduction.

·         Only organisms that reproduce asexually can produce offspring that are exact copies of themselves.

·         In asexual reproduction, a single individual is the sole parent to donate genes to its offspring.

°         Single-celled eukaryotes can reproduce asexually by mitotic cell division to produce two genetically identical daughter cells.

°         Some multicellular eukaryotes, like Hydra, can reproduce by budding, producing a mass of cells by mitosis.

·         An individual that reproduces asexually gives rise to a clone, a group of genetically identical individuals.

°         Members of a clone may be genetically different as a result of mutation.

·         In sexual reproduction, two parents produce offspring that have unique combinations of genes inherited from the two parents.

·         Unlike a clone, offspring produced by sexual reproduction vary genetically from their siblings and their parents.

B. The Role of Meiosis in Sexual Life Cycles

·         A life cycle is the generation-to-generation sequence of stages in the reproductive history of an organism.

°         It starts at the conception of an organism and continues until the organism produces its own offspring.

1. Human cells contain sets of chromosomes.

·         In humans, each somatic cell (all cells other than sperm or ovum) has 46 chromosomes.

°         Each chromosome can be distinguished by size, position of the centromere, and pattern of staining with certain dyes.

·         Images of the 46 human chromosomes can be arranged in pairs in order of size to produce a karyotype display.

°         The two chromosomes comprising a pair have the same length, centromere position, and staining pattern.

°         These homologous chromosome pairs carry genes that control the same inherited characters.

·         Two distinct sex chromosomes, the X and the Y, are an exception to the general pattern of homologous chromosomes in human somatic cells.

·         The other 22 pairs are called autosomes.

·         The pattern of inheritance of the sex chromosomes determines an individual’s sex.

°         Human females have a homologous pair of X chromosomes (XX).

°         Human males have an X and a Y chromosome (XY).

·         Only small parts of the X and Y are homologous.

°         Most of the genes carried on the X chromosome do not have counterparts on the tiny Y.

°         The Y chromosome also has genes not present on the X.

·         The occurrence of homologous pairs of chromosomes is a consequence of sexual reproduction.

·         We inherit one chromosome of each homologous pair from each parent.

°         The 46 chromosomes in each somatic cell are two sets of 23, a maternal set (from your mother) and a paternal set (from your father).

·         The number of chromosomes in a single set is represented by n.

·         Any cell with two sets of chromosomes is called a diploid cell and has a diploid number of chromosomes, abbreviated as 2n.

·         Sperm cells or ova (gametes) have only one set of chromosomes—22 autosomes and an X (in an ovum) and 22 autosomes and an X or a Y (in a sperm cell).

·         A gamete with a single chromosome set is haploid, abbreviated as n.

·         Any sexually reproducing species has a characteristic haploid and diploid number of chromosomes.

°         For humans, the haploid number of chromosomes is 23 (n = 23), and the diploid number is 46 (2n = 46).

2. Let’s discuss the role of meiosis in the human life cycle.

·         The human life cycle begins when a haploid sperm cell fuses with a haploid ovum.

·         These cells fuse (syngamy), resulting in fertilization.

·         The fertilized egg (zygote) is diploid because it contains two haploid sets of chromosomes bearing genes from the maternal and paternal family lines.

·         As an organism develops from a zygote to a sexually mature adult, mitosis generates all the somatic cells of the body.

°         Each somatic cell contains a full diploid set of chromosomes.

·         Gametes, which develop in the gonads (testes or ovaries), are not produced by mitosis.

°         If gametes were produced by mitosis, the fusion of gametes would produce offspring with four sets of chromosomes after one generation, eight after a second, and so on.

·         Instead, gametes undergo the process of meiosis in which the chromosome number is halved.

°         Human sperm or ova have a haploid set of 23 different chromosomes, one from each homologous pair.

·         Fertilization restores the diploid condition by combining two haploid sets of chromosomes.

3. Organisms display a variety of sexual life cycles.

·         Fertilization and meiosis alternate in all sexual life cycles.

·         However, the timing of meiosis and fertilization does vary among species.

·         These variations can be grouped into three main types of life cycles.

·         In most animals, including humans, gametes are the only haploid cells.

°         Gametes do not divide but fuse to form a diploid zygote that divides by mitosis to produce a multicellular organism.

·         Plants and some algae have a second type of life cycle called alternation of generations.

°         This life cycle includes two multicellular stages, one haploid and one diploid.

°         The multicellular diploid stage is called the sporophyte.

°         Meiosis in the sporophyte produces haploid spores that develop by mitosis into the haploid gametophyte stage.

°         Gametes produced via mitosis by the gametophyte fuse to form the zygote, which grows into the sporophyte by mitosis.

·         Most fungi and some protists have a third type of life cycle.

°         Gametes fuse to form a zygote, which is the only diploid phase.

°         The zygote undergoes meiosis to produce haploid cells.

°         These haploid cells grow by mitosis to form the haploid multicellular adult organism.

°         The haploid adult produces gametes by mitosis.

·         Note that either haploid or diploid cells can divide by mitosis, depending on the type of life cycle. However, only diploid cells can undergo meiosis.

·         Although the three types of sexual life cycles differ in the timing of meiosis and fertilization, they share a fundamental feature: each cycle of chromosome halving and doubling contributes to genetic variation among offspring.

4. Meiosis reduces the chromosome number from diploid to haploid.

·         Many steps of meiosis resemble steps in mitosis.

°         Both are preceded by the replication of chromosomes.

·         However, in meiosis, there are two consecutive cell divisions, meiosis I and meiosis II, resulting in four daughter cells.

°         The first division, meiosis I, separates homologous chromosomes.

°         The second, meiosis II, separates sister chromatids.

·         The four daughter cells have only half as many chromosomes as the parent cell.

·         Meiosis I is preceded by interphase, in which the chromosomes are replicated to form sister chromatids.

°         These are genetically identical and joined at the centromere.

°         The single centrosome is replicated, forming two centrosomes.

·         Division in meiosis I occurs in four phases: prophase I, metaphase I, anaphase I, and telophase I.

Prophase I

·         Prophase I typically occupies more than 90% of the time required for meiosis.

·         During prophase I, the chromosomes begin to condense.

·         Homologous chromosomes loosely pair up along their length, precisely aligned gene for gene.

°         In crossing over, DNA molecules in nonsister chromatids break at corresponding places and then rejoin the other chromatid.

°         In synapsis, a protein structure called the synaptonemal complex forms between homologues, holding them tightly together along their length.

°         As the synaptonemal complex disassembles in late prophase, each chromosome pair becomes visible as a tetrad, or group of four chromatids.

°         Each tetrad has one or more chiasmata, sites where the chromatids of homologous chromosomes have crossed and segments of the chromatids have been traded.

°         Spindle microtubules form from the centrosomes, which have moved to the poles.

°         The breakdown of the nuclear envelope and nucleoli take place.

°         Kinetochores of each homologue attach to microtubules from one of the poles.

Metaphase I

·         At metaphase I, the tetrads are all arranged at the metaphase plate, with one chromosome facing each pole.

°         Microtubules from one pole are attached to the kinetochore of one chromosome of each tetrad, while those from the other pole are attached to the other.

Anaphase I

·         In anaphase I, the homologous chromosomes separate. One chromosome moves toward each pole, guided by the spindle apparatus.

·         Sister chromatids remain attached at the centromere and move as a single unit toward the pole.

Telophase I and cytokinesis

·         In telophase I, movement of homologous chromosomes continues until there is a haploid set at each pole.

°         Each chromosome consists of two sister chromatids.

·         Cytokinesis usually occurs simultaneously, by the same mechanisms as mitosis.

°         In animal cells, a cleavage furrow forms. In plant cells, a cell plate forms.

·         No chromosome replication occurs between the end of meiosis I and the beginning of meiosis II, as the chromosomes are already replicated.

Meiosis II

·         Meiosis II is very similar to mitosis.

°         During prophase II, a spindle apparatus forms and attaches to kinetochores of each sister chromatid.

§         Spindle fibers from one pole attach to the kinetochore of one sister chromatid, and those of the other pole attach to kinetochore of the other sister chromatid.

·         At metaphase II, the sister chromatids are arranged at the metaphase plate.

°         Because of crossing over in meiosis I, the two sister chromatids of each chromosome are no longer genetically identical.

°         The kinetochores of sister chromatids attach to microtubules extending from opposite poles.

·         At anaphase II, the centomeres of sister chromatids separate and two newly individual chromosomes travel toward opposite poles.

·         In telophase II, the chromosomes arrive at opposite poles.

°         Nuclei form around the chromosomes, which begin expanding, and cytokinesis separates the cytoplasm.

·         At the end of meiosis, there are four haploid daughter cells.

5. There are key differences between mitosis and meiosis.

·         Mitosis and meiosis have several key differences.

°         The chromosome number is reduced from diploid to haploid in meiosis but is conserved in mitosis.

°         Mitosis produces daughter cells that are genetically identical to the parent and to each other.

°         Meiosis produces cells that are genetically distinct from the parent cell and from each other.

·         Three events, unique to meiosis, occur during the first division cycle.

1.       During prophase I of meiosis, replicated homologous chromosomes line up and become physically connected along their lengths by a zipperlike protein complex, the synaptonemal complex, in a process called synapsis. Genetic rearrangement between nonsister chromatids called crossing over also occurs. Once the synaptonemal complex is disassembled, the joined homologous chromosomes are visible as a tetrad. X-shaped regions called chiasmata are visible as the physical manifestation of crossing over. Synapsis and crossing over do not occur in mitosis.

2.       At metaphase I of meiosis, homologous pairs of chromosomes align along the metaphase plate. In mitosis, individual replicated chromosomes line up along the metaphase plate.

3.       At anaphase I of meiosis, it is homologous chromosomes, not sister chromatids, that separate and are carried to opposite poles of the cell. Sister chromatids of each replicated chromosome remain attached. In mitosis, sister chromatids separate to become individual chromosomes.

·         Meiosis I is called the reductional division because it halves the number of chromosome sets per cell—a reduction from the diploid to the haploid state.

·         The sister chromatids separate during the second meiosis division, meiosis II.

C. Origins of Genetic Variation

·         What is the origin of genetic variation?

·         Mutations are the original source of genetic diversity.

·         Once different versions of genes arise through mutation, reshuffling during meiosis and fertilization produce offspring with their own unique set of traits.

1. Sexual life cycles produce genetic variation among offspring.

·         The behavior of chromosomes during meiosis and fertilization is responsible for most of the variation that arises in each generation.

·         Three mechanisms contribute to genetic variation:

1.       Independent assortment of chromosomes.

2.       Crossing over.

3.       Random fertilization.

·         Independent assortment of chromosomes contributes to genetic variability due to the random orientation of homologous pairs of chromosomes at the metaphase plate during meiosis I.

°         There is a fifty-fifty chance that a particular daughter cell of meiosis I will get the maternal chromosome of a certain homologous pair and a fifty-fifty chance that it will receive the paternal chromosome.

·         Each homologous pair of chromosomes segregates independently of the other homologous pairs during metaphase I.

·         Therefore, the first meiotic division results in independent assortment of maternal and paternal chromosomes into daughter cells.

·         The number of combinations possible when chromosomes assort independently into gametes is 2n, where n is the haploid number of the organism.

°         If n = 3, there are 23 = 8 possible combinations.

°         For humans with n = 23, there are 223, or more than 8 million possible combinations of chromosomes.

·         Crossing over produces recombinant chromosomes, which combine genes inherited from each parent.

·         Crossing over begins very early in prophase I as homologous chromosomes pair up gene by gene.

·         In crossing over, homologous portions of two nonsister chromatids trade places.

°         For humans, this occurs an average of one to three times per chromosome pair.

·         Recent research suggests that, in some organisms, crossing over may be essential for synapsis and the proper assortment of chromosomes in meiosis I.

·         Crossing over, by combining DNA inherited from two parents into a single chromosome, is an important source of genetic variation.

·         At metaphase II, nonidentical sister chromatids sort independently from one another, increasing by even more the number of genetic types of daughter cells that are formed by meiosis.

·         The random nature of fertilization adds to the genetic variation arising from meiosis.

·         Any sperm can fuse with any egg.

°         The ovum is one of more than 8 million possible chromosome combinations.

°         The successful sperm is one of more than 8 million possibilities.

°         The resulting zygote could contain any one of more than 70 trillion possible combinations of chromosomes.

°         Crossing over adds even more variation to this.

·         Each zygote has a unique genetic identity.

·         The three sources of genetic variability in a sexually reproducing organism are:

1.       Independent assortment of homologous chromosomes during meiosis I and of nonidentical sister chromatids during meiosis II.

2.       Crossing over between homologous chromosomes during prophase I.

3.       Random fertilization of an ovum by a sperm.

·         All three mechanisms reshuffle the various genes carried by individual members of a population.

2. Evolutionary adaptation depends on a population’s genetic variation.

·         Darwin recognized the importance of genetic variation in evolution.

°         A population evolves through the differential reproductive success of its variant members.

°         Those individuals best suited to the local environment leave the most offspring, transmitting their genes in the process.

·         This natural selection results in adaptation, the accumulation of favorable genetic variations.

·         If the environment changes or a population moves to a new environment, new genetic combinations that work best in the new conditions will produce more offspring, and these genes will increase.

°         The formerly favored genes will decrease.

·         Sex and mutation continually generate new genetic variability.

·         Although Darwin realized that heritable variation makes evolution possible, he did not have a theory of inheritance.

·         Gregor Mendel, a contemporary of Darwin ’s, published a theory of inheritance that supported Darwin ’s theory.

°         However, this work was largely unknown until 1900, after Darwin and Mendel had both been dead for more than 15 years.