Selection occurs within a population but evolution requires generations.
show evolution over long time frames
artifical selection
yes
1. variation is pre existing and can be passed on to offspring
2. selection is based on environment
3. not all offspring survive (those who survive is non random)
4. differential reproduction (those with advantageous traits in that space and time are more likely to survive and reproduce)
didn't propose a theory for inheritance, the concept of mutation was not developed until 1900's
transformational evolution
variational evolution
fossils
an environment where they can fossilize
george cuvier first started studying fossils and first proposed extinctions
it is incomplete, we've only found the easiest to find fossils, hot humid areas rarely have fossils. fossils are also biased because hard and bony parts/ aquatic organisms fossilize better.
due to changing sediment conditons, if the environment that the fossil was in changes it can obscure the date
universe- 13.77 byo and earth- 4.5 byo
because of 4.4 byo meteorite in australia and zircon crystals that are also this old, darwin also hypothesized this because of geological formations in england
at a measurable rate
at dawn of solar system
that our solar system is ancient
approx 3.8byo, esitmated by principle of superpostion
520 Mya: fossils of oldest known chordates (incl vertebrates)
480 Mya: the oldest trackways
475 Mya: oldest land plant fossils
360 Mya: oldest land vertebrates
230 Mya: dinosaurs
180 Mya: Our common mammalian ancestor
150 Mya: birds
no just zircons and stromatolites
deep sea vents and hot springs
that there was tons of energy, carbon and lots of redox worthy chemicals
that there was tons of energy and nutrients, lots of oxidized minerals
where they tried to recreate the conditons of early earth but failed
very little oxygen, lots off ammonia, lots of methane, lots of volcanic activity. These conditions together could cause abiotic synthesis of organic molecules
1.8bya because of acritarchs
used to think 1.6 byo, interpreted as filamentous algae
sponges
known as the ediacaran fauna
segmentation, 3 axis's, more complexity. (ex. trilobite, mollsucs, and first chordates)
Increased O2, allowed more primary productivity, allows emergence of predators, allows inc in body size BUT oxygen levels didn’t necessarily change around Cambrian and larger animals already existed
Evolutionary innovation, evolution of novel traits (ex. Segmentation) allowed increased adaptability (ex. Hox gene cluster evolution may have sparked Cambrian)
Predation and arms race, if predation arose during Cambrian it could explain, inc body size, hard body size, motility/ swimming ability
the late Ordovician and early Silurian period (450 Mya). Most likely liverwort like organisms, possibly co invaded by fungi, vascular plants evolved after
millipede
amniotes produce shelled eggs
this was the paleogene mass extinction event which caused the loss of big reptiles but the little mammals thrived (prob due to loss of predators)
Darwin thought, more shared features= more recent common ancestor
vary with the taxa of interest, have different states, can be morphological, behavioural, biochemical, genetic.
to reconstruct evolutionary relationships so that trees can reflect shared ancestry
life, domain, kingdom, phylum, class, order, family, genus, species
mendel determined that traits of offsprings were determined by parental factors that remained discrete.
traits that have no blending
hand pollinated pea plants with known traits, he was the first to use math to explain predicted traits in offspring
Crossed round and wrinkled peas, all progeny were round (F1)- all showed only one parental phenotype, no blend, these were heterozygote (het) and showed the dominant traits. Then he crossed Rr and Rr and got progeny that were 3 R: 1 W (approximately).
that traits came in pairs and the genes are halved in germ cells
law of dominance, law of segregation, law of independent assortment
most traits dont follow the phenotypic ratios of mendelian inheritance because of multiple alleles of one gene, influence on other genes and influence of environment
a diploid (2n) will only carry 2 alleles in an individual but multiple alleles can exist in population which can create variation in a population without necessarily being influenced by other genes or the environment
ex. pavilion dardanus butterlfies which have multiple alleles which give rise to multiple wing patterns within the same speices
ex. human ABO blood type has 3 alleles, 9 possible genotype within pop, 4 possible phenotypes
most traits are continous
where their phenotypic variation exists on spectrum
polygenic inheritance, where many genes contribute to the same traits
environmental conditions can have a great impact on phenotype, even greater than genotype sometimes.
These conditions can include, temp, availability of nutrients, presence of toxins/mutagen, some can be the maternal effect
Easiest way is to force mate individuals with opposite phenotypes if the progeny will follow a Mendelian inheritance pattern in the F2 but most traits aren’t Mendelian.
it is arranged into chromosomes, multiple linear chrosomes. DNA in eurkaryotes is organzied using histone proteins. The DNA wraps around a histone octomer creating a nucleosome.
The organization of DNA protects it from damage, organizes the nucleus and limits access of transcriptional machinery to DNA
chromsomes are stored in the eukaryotic nucleus (membrane bound organelle), chromosome sizes can vary greatly- msot eukaryotes are diploid (2n)
in the soma of humans cells are diploid but germ cerlls are haploid
DNA to RNA (transcription), RNA to protein (translation). Only one strand of DNA is used as a template.
Translation is carried out by ribosome and uses a universal codon table. 3 nucleotides= 1 amino acid
Redundancy in the genetic code allows mutational tolerance (more than one codon encoding same amino acid. Most of redundancy is @ 3rd codon.)
In most euk, each cell contains the exact same entire genome. Only a fraction of genes are expressed giving rise to different cell types.
Most gene regulation occurs at the level of transcription initiation. This is regulated by a region of the DNA called the promoter.
alternative splicing
Most of eukaryotic genome is not expressed and most doesn’t become protein
genes that have mutated such that they no longer contain a promoter, hence not being transcribed to mrna
markers are specific sequences of DNA, often located throughout the genome, can be used as refrerence points to track the inheritance of genes linked to certain traits. the goal is to find genetic markers significantly associated with the extremes of the trait
jumping genes, products of ancient viral infection. Can change positions within the DNA, can cause mutations by disrupting the genes or regulatory regions that they jump to.
no gemone size is unrelated to complexity, genome size caries greatly between organisms. ex. sea anemones have more genes than humans.
from alternative splicing not from gene #
process by which a single gene can produce multiple different versions of protein, occurs during RNA processing. alternative splicing allows for the exons (coding regions) to be arrange in different combinations leading to the production of multiple proteins from the same gene.
meiosis
4 genetically different haploid gametes
crossing over, independent assortment, random fertilization
pair of chromsomes have same gene order but potentially different alleles
it generate genetic variation and haploid gametes.
due to the crossing over in prophase I (homo chromo exchange genetic material)
and
independent assortment in metaphase I (homo chromo pairs line up, arrange randomly)
and
random fertilization (random sperm and random egg)
only the environment determines if a mutation is harmful (most common), neutral, beneficial (rare).
they rely on very rare benefical mutations for variation
silent muation, missense mutation, nonsense mutation, frameshift mutation
change in a single base pair of DNA sometimes due to DNA replication error or exposure to mutagens
change in DNA seq that doesnt change protein seq
ex. GAA changed to GAG but both of these code for the same amino acid
a change in a single base pair that results in a codon coding for a different amino acid which can affect protein function
ex. GAG mutated to GTG code for a different amino acid which can affect the protein.
point mutation that changes codon to a stop codon, causing protein to be prematurely shortened. Can cause to loos of function or non fuctional proteins.
ex. CAA muated to UAA (stop codon), causes translation to stop early.
deletetion/insertion of nucleotide so that the reading frame shifts of the genetic sequence.this leads to a complelety different protein sequence after the point of mutation which can be devastating to protein function
missense maybe, nonsense, frameshift
no they are common, but the rarer events have greater effects
yes, gain/loss of chromosome
gain or loss of chromosomes, often have great effects
yes
around 61
HW allows us to predict the particular phenotypes in offspring for an entire pop
HW eqbrm provides a null hypothesis for testing evidence of evolutionary change.
if there was no selection/mutation/drift
no, to be satisifed a population must be infinitely large, have no net muation, no net migration, mate randomly, and have no selection (no disadvantageous/advantageous traits)
alleles and genotype freq in offspring would be the same as parent gen
P^2 + 2pq + q^2= 1
p= dominant allele
q= recessive allele
a1/a1=0.25, a2/a2=0.25, a1/a2=0.50
p=0.6 and q=0.4. P+Q=1
refer to the proportions of different genotypes (homozygous domiannt, heterozygous, homo recessive) in a pop.
genotypic frequency can shift due to non random mating or genetic drift. allele freq shift from mirgation/mutations
no
p= 0.7625
q=0.2375
p^2= 0.581 (MM)
2pq= 0.362(MN)
q^2= 0.0564 (NN)
MM 0.58 of 320= 185
MN 0.36 of 320= 115
NN 0.05 of 320= 16
roughly match
the connection between the pop
evolution- mutation, natural selection, migration,drift
no, sometimes random events like drift and mutation happen
because it is random and no selected for.
computer simulation or with a drosphila lab experiment
impact of drift is dependent on pop size. The larger the population the more genetic stability. The smallest populations have the biggest fluctuations and fixation is least likely
researcher mates 8 females to 8 males each gen, 107 lab pop of D.melanogaster each found with 16 bw/bw^75 heterozygotes. Found: over 19 gens, bw^75 allele is lost, bw allele is fixed after 30 gen.
when every indiv in the pop has the same allele at a particular gene location, no genetic variation for that particular gene in the pop- only one allele exists and it is the only form of that gene being passed on
yes, any small shifts in allele freq plugged into HW would turn into big shifts in genotype freq.
big problem for conservation because don't necessarily want the 2 extremes and drift makes the wild extreme more common. At a point of a species being endangered the odds of an allele being fixed increases
bottleneck and founder effect.
related to how common it is in the original population. Rarer alleles simply less likely to make through bottleneck, more likely to be lost. Ex: northern elephant seal. Was nearly hunted to extinction (20 indiv left), population expansion after a constriction lead to the bottleneck effect now observable in the genetic diversity of the extant population
small group of indiv from a larger pop start a new pop, the founders are the only subset of genetic diveristy present. As a result, new pop may have different allele freq than original pop and certain alleles may be over or underrepresented
ex. the mutiny of the bounty
the calculated egree to which an indiv is inbred. F= proportion of individuals in a population who are autozygous.
allele that is inherited identically by descent from both parent, allele comes from common ancestor. both copies of allele are identical
two copies of a gene in an indivual are not identical by descent, no common ancestor. 1-F= fraction that are allozygous
if no in breeding, F=0, het= 2pq= HW
If there is inbreeding, F>0, F<1, the proportion of 2pq= less than H-W predictions.
F=0 not in bred for gold F=0.5 inbred for purple
inbreeding does not change allele freq but increases homozygosity
allows recessive, deleterious alleles to persists and reduce viability. Their effect only obvious when homozygous.
the hapsburgs family where 50% hapsburgs children died before during 1 where as the rest of spanish pop ~ 80% survived
when populations are separated by geographical barriers, because members of pop tend to be sedentary then gene flow is low, then pop will genetically differentiate over time due to drift.
lynx have greater home ranges so over the same distances show greater connectdetness than big horn sheep
usually, this is bc that means there is been a greater amount of time that has passed since the two pops have interacted. These distances can also be used as pop fingerprints. To determine where an indiv came from
Landscape features (including man made) can influence home ranges and gene flow.
More gene flow means increased variation means less drift.
first filial generation
movement of alleles between different pops through migration or interbreeding of indiv. increase genetic diversity within a pop by introducing new alles from another pop, it can reduce genetic differences between populations, and it can prevent speciation (formation of new species) by maintain connectivity.
knowledge of geography allows researchers to explore how genotypes vary across landscapes.
(type of drift) when a population undergoes a rapid constriction in population size, then expands = the allele freq in surviving population may not reflect the freq in original population
(type of drift), occurs when an offshoot of a bigger population establishes a new population, therefore the new population will be dependent on alleles the founders carry. Typically the alleles that are most common will be most represented (but there is randomness)
inbreeding increases homozygosity and brings recessive (and sometimes deleterious) alleles to surface. Declines in survival and fecundity are due to inbreeding humans.
mating with close relatives, violates HW assumption of random mating. Selfing is the most extreme form, and shows how inbreeding reduces the proportion of heterozygotes in a population.
suppose we could magically label individual alleles and follow them through subsequent generations. Ex. Label one copy of A1 in a male with an asterisk (*). Potential for diversity is reduced. after 1 gen A1* may have been passed to both son and daughter, if siblings mate= 25% chance that offspring will have both alleles from grandfather
individuals can interact with one another very easily (pigeons)
there are little pockets of individuals that can move between subdivision but its not contiguous (snowshoe hares in edm)
happens when the H-W assumption of an infinite population is violated. Random changes in the frequency of two or more alleles/haplotypes/genotypes therefore we have to consider drift as a force that can cause allelic change (evolution) that are not due to selection.
populations that are completely isolated from one another
group of interacting (potential) and potentially interbreeding individuals of a particular species
the study of allele distributions within and among populations and how they change over time
different forms arise from changes in allele frequency. it is specifically defined as a change in inherited traits in a population over time, Darwin described it as descent with modification. Evolution is not synonymous with natural selection, natural selection is a mechanism of evolution but not the only mechanism and it may result in evolution if selection causes a change in allele frequencies (genetic structure of pop) over time.
muation, drift, natural select, gene flow.
new: aritifcal selection
mechanism that drives heritable variation (not all mutations do this)
random changes to the genetic makeup of a population (mutation is a mechanism of drift, there are several others too)
caused by the interaction of genotype with the environment.
migration between populations
now an important driver of evolution (goes hand in hand with natural selection, new)
Rate of reproduction
Selective forces
Environmental change
anatomist, found life forms that went extinct. Thought extinction was only caused by catastrophe
zoologist, curated fossils and extant mollusks. Saw gradual change in individuals. Suggested species change over time based on the use and disuse theories. Also that change comes from interactions with the environment that can be passed onto offspring.
He got a few things wrong ( that microbes arise from spontaneous generation, that humans were the most complex species therefore they have been around the longest and the inheritance of acquired traits which is partially wrong because it has some applications.)
geologist, found that landforms are not fixed but slowly changing because of geological processes, estimated earth was older than biblical 6000 years. Also proposed extinction.
talked about survival of the fittest in a human population. Human pop increases faster than food supply (demand outpaces supply). Completion leads to survival of the fittest because resources are always limited .
Naturalist, came up with natural selection independent of Darwin. Encouraged Darwin to publish.
Collected specimens and fossils found patterns of distribution of traits (particularly distribution of beak lengths in finches between Galápagos Islands). Wrote “ the origin of the species by natural selection). Descent with modification but he did not understand how modification or inheritance occurred.
far from mainland South America, relatively young islands, little gene flow isolated pop were a result of bottle neck.
Variation- exists in all species
Artificial selection- enhancement of desired traits
Distribution of species in time and space- if there is descent with modifications, then similar species with modifications should be found in similar areas.
Transformational evolution, forms change in their lifetime and that is passed on . Everything changes and change is heritable
Variational evolution, only a few survive and persist
trace of past life that has fossilized through a slow process of mineralization, rare but numerous, many organisms don’t fossilize because it required perfect conditions of desiccation (dehydration) and mineralization. Soft tissues rarely fossilize.
bones/ teeth- readily fossilize
indirect evidence of life, not of the organism but from the organism (ex. Subfossil: fossilized footprints, a coprolite (poop). Important for determining the physiological method of the organism.
organisms engage in chemical reactions that leave a mark in geological time (ex. Fossils fuels represent carbonaceous deposits of fossilized marine algae)
crystals the preserve specks of carbon used for dating
based on the principle of radioactive decay, these radioactive isotopes decay at a steady state (denoted by their half life)
amount of time for half of the atoms in a unit of element to decay to daughter products
largest unit of time (can be thousands or millions of years long), delineated by major changes to earth's system
no evidence of life, very volatile, inhospitable, no liquid water
earliest forms of unicellular life, life was limited to the oceans ( which contained oxygen, nutrients and energy) as with only trace oxygen in the atmosphere. Bacteria first then Archaea. stromatolites.
dating the rock layering to date fossilized chemical life forms in that layer (estimate). Uses radiometric dating (ex. Uranium for old things and carbon for young things (<50k years).
allows us to date tissue itself, can be naturally replenished in atmosphere (< 50k years)
layered structures formed by chemical reactions of ancient bacteria (oldest fossils). Ex. In Ontario there are iron banded formations - chemical signatures of ancient life, only from biotic processes.
beginnings of eukaryotic and multicellular life. Great oxygenation Event (2.5 bya- 2.2 bya), oxygen content in the atmosphere increased significantly due to the rise of photosynthetic bacteria. Oxygen is a strong selective pressure because it is so reactive, this along with other pressures probably led to the evolution of eukaryotes.
the Cambrian explosion marked the evolution of the modern day animal body plan, this eon and forwards is depicted in greater detail, more visible life.
Paleozoic (old life), Mesozoic (middle life), Cenozoic (new life)
LUCA: last universal common ancestor
Prokaryotes: no membrane bound nucleus (bacteria and archaea)
Eukaryotes: archaea evolved internal membranes
Archaea: “extremophile”, prokaryotic, ancient archaeon that evolved in eukaryote.
single celled organisms of ambiguous origin. Large, structurally complex biological specimens. How we presumed origin of eukaryotes .
has this biomarker (chemical signature)- cholesterol like molecules
mostly fossilized sponges, jellyfish and comb jellies, front like organisms. Small animals with simple morphology. Almost completely wiped out by a mass extinction.
most extensive period of adaptive radiation and formation of new life forms, around 541 Mya the entire ocean ecosystem was reorganized and new animal body plans emerged, Cambrian animals had modern body plans (invention of segmentation and 3 axis (r/l, d/v, a/p), notochord, more complex). Most major extant animal phyla make their first appearance in deposits (550 Mya) . Appearance of hard body parts, and huge variety of form, feeding methods and locomotion. Best known deposit is the Burgess Shale.
vascular plants evolved. Earth was covered in ferns, horsetails and first seed plants . Land plants got more sun but they need to stand up and were susceptible to dehydration
produced shelled eggs. 3 forms - diapsids (2 holes behind eye sockets like reptiles), synapsids (1 hole behind eye sockets like horses), and dinosaurs (230 Mya- Mesozoic).
age of mammals (66 Mya-present), earliest fossils of ape ancestry. 55 Mya- primates, 20 Mya- apes, 7 Mya- hominins, 0.2 Mya- Homo sapiens
evolutionary history of a lineage
Visual representation of phylogeny
developed the first real trees, included time scales, trees can demonstrate evolutionary relationship and time.
worked on fly taxonomy and came up with an approach to phylogenetics that was more systematic; cladistics.
relationships among taxa follow a dichotomous branching pattern= branch in pairs
Shared, derived traits (synapomorphies) that provide for relative recentness in common ancestry
Principle of parsimony
Cladogram should be consistent with the inferred pattern of historical relatedness
A species known to have or suspected to have split off prior to main diversification event, are necessary to root the tree (a distinct starting point). The outgroup is not a primitive species but it does have ancestral traits and other derived traits
0: in outgroup considered ancestral
1: a state that is different than outgroup
shared, derived trait. Trait that is present in more than one group. found in 2 or more speices from common ancestor. trait that is new to the clade and distinguishes that clade from other organisms.
shared, ancestral trait. Present in both Ingroup and outgroup
ex. vertebral colum for vertebrates.
derived trait found in only one taxon. only in one species. defines a single lineage
ex. long neck for giraffes even within mammals unique for giraffe
represents convergent evolution, character states that are present in more than one taxon but arose independently due to similar environmental pressures or functional needs
winga of bats, birds, insects
character states that are present in more than one taxon but arose from a common ancestor
limbs of humans, cats, whales, bats
comb, represents an unresolved tree
scientific study of all kinds and diversity of organisms and all relationships among them
science of describing, naming, and classifying species of living and fossilized organisms. (Is subject to change)
the study of evolutionary relationships among organisms
created the first systematic binomial naming system for organisms. Each organisms was given a genus, specific epithet (species name). Created the 3 kingdoms, plants (regnum vegtabile), animals (regnum animals), minerals (regnum Lapideum). Then the smaller categories, class, order,genus, species. His mistakes was that minerals aren’t alive and that humans as different subspecies, also didn’t believe species could evolve
No two species within the same kingdom can have the same name
You can’t name stuff after yourself but you can name it after other people.
Has to grammatically make sense in Latin, but sometimes using indigenous names from the region the specimens are found (ex.tiktaalik roseae- first animals to leave ocean)
Classify organism by domain, kingdom, phylum, subphylum, class, order, family, genus and species
Only genus and species are italicized/ underlined.
Everything except species should be capitalized
Each successive level is nested within another
designated example specimen(s) representing a species. Used to determine if an unknown organism belongs to that species. Usually considered the best representation of a species, housed in museums and collections, they must be available to the public and to researchers
variants of a gene
single designated specimen used a ‘type’, the ‘name bearer’.
collection of specimens used a type for a species
the genetic makeup of an organism, genotypes are heritable. Individuals inherit alleles. One genotype can lead to several different phenotypes depending on how the gene is expressed (genotype + environment = phenotype).
the physical manifestation of that sequence is the phenotype, the phenotypes are not inherited. The phenotype (manifestation of the gene) is what selection acts on, can include physiological processes, developmental processes, behaviours
the dominant allele is expressed in a het (heterozygote).
all alleles are equally represented in gametes. 50;50 chance of giving one allele to offspring.
genes on different chromosomes assort independently (ex. Colour and shape are independently assorted/inherited).
more than 1 discrete trait can arise from a single genotype at a single locus (genetic location= gene), due to the influence of environment. Ex. Aphids can be wingless or winged as adults with the same genotype because they have different phenotypes from nutrition profiles during development. This is an irreversible phenotype.
study of genetics of extremes- look for variants/loci that are more likely associated with the extreme traits. Sometimes the loci found are not the genes themselves but are genetically linked to the determinant locus.
the log of odds, ex. lod score of 2 means 100x likelihood of being associated with height.
the same genotype can manifest as different phenotypes depending on the environment and can change in a lifetime. Ex. Hydrangea flower color can change based on soil pH. If soil has neutral ph- flower is white, if soil is acidic- flower is blue, if solid is basic-flowers are pink. You can take white flower cuttings and grow in soil of different ph and the colour changes, same genotype but different phenotype.
Take two individuals with distinct phenotypes and environments and grow them in the same environment to see if those phenotypes stay. Ex. Blue flower plants tend to grow on east slopes, white flower plants tend to grow on west slopes and then we swap them, after moving to a new environment the plants display a parental phenotype therefore it is most likely genetic.
the machines of the cell and where variation allows selection. chains of amino acids folded into unique 3D structures, structures can be simple or complex. Structures can be made from one amino acid chain= monomer. Structures can be made from multiple amino acids= oligo or polymer.
a polymer of nucleotides arranged into a double helix, it is double stranded, antiparallel (strands of DNA run in opposite directions of each other), complementary ( A=T, G=C).
2 genes that make one functional protein.
a sequence of DNA that encodes a trait that can be selected for.
molecules of DNA where there are thousands of genes.
a sequence of DNA that regulates transcription initiation. Bound by proteins called transcription factors.
generates genetically identical daughter cells (diploid to diploid)
generates genetic variation and haploid gametes (diploid to haploid)
physical exchange of DNA between pairs of chromosomes.
2 pairs of chromosomes assort independently (equally likely segregation patterns)
sperm that fertilizes isn;t necessarily the strongest, all sperm are different (and eggs).
change to the genetic sequence or content of an organism, they are common and occur during DNA replication (whenever DNA is being copied there is a chance of error). Many mutagens (things that cause mutations) in the environment like UV light.
Every mutation is equally likely therefore every nucleotide can change. Mutations are random/non directional, and comes before selection, environment determines if the mutation is beneficial, neutral or harmful (not adaptively directed).
Most mutations are not inherited and they must occur in germline to be heritable, most mutations happen in soma.
substitution of a single base with a different base, can be transitions of transversions. Has to happen at the same position on the same strand. In DNA a change in sequence is reflected in mRNA. If a point mutation happens in a coding region it can result in a simple protein mutation, can be synonymous or no synonymous.
purine with purine (A-G) or pyrimidine (C-T) with pyrimidine
pyrimidine with purine or vice versa
silent, no effect on amino acid sequence= mutational tolerance due to redundancy in the genetic code.
change amino acid sequence, can be missense (encodes the wrong aa and may affect the structure/function of a protein), can be non sense (creates a premature stop codon and a truncated protein, this will affect the structure and may affect the function)
these are insertions and deletion of 1+ nucleotides. If indels occur in a protein encoding sequence it causes a frame shift that affects all amino acids/codons after index.
flipping of a DNA segment within a chromosome, often causes no major changes.
repeating the same sequence often in tandem with the original ex. ABCD—> ABCBCD (can have dosage effects, often a result of transposable elements- jumping genes)
originally separate chromosomes to break and fuse with another chromosome
the #, size, shape, arrangements of an organism's chromosomes. Change in karyotype can affect fertility of an individual and viability of the offspring.
unbalanced # of chromosomes, 2n +1 or 2n-1
duplication of entire genome, often results in sterility (esp if off # of chromsomes
4n, are more fertile than triploids (3n)
genome duplication within same species
genome amplification due to hybridization between 2 species