BIOL 230 Midterm I, 2/12/15 Name_____________________________________

Biology  230  
2015:  Midterm  I  
 
This  is  a  closed  book/notes  exam;  use  of  calculators  is  fine.    Please  write  in  ink  so  that  we  may  
regrade  upon  request.    You  will  have  the  entire  period  in  which  to  complete  the  exam.    A  list  of  
formulae  from  population  and  quantitative  genetics  is  provided  on  the  last  page.    Please  write  
your  name  on  ALL  pages  of  the  exam,  as  pages  may  be  separated  to  facilitate  grading.    Please  
show  all  your  work  so  that  we  may  give  partial  credit.  
 
1.  Clearly  (but  succinctly)  define  and  state  the  significance  of  the  following  terms  (5pts.  each).  
 
adaptation  
 
 
 
 
 
 
 
 
heritability  
 
 
 
 
 
 
 
 
 
2.  How  does  Darwin  frame  natural  selection  in  terms  of  the  struggle  for  existence  and  adaptation?  
How  are  population  and  food  related  in  Darwin's  view  of  the  struggle  for  existence,  as  derived  
from  Malthus?  (5pts)  
 
 
 
 
 
 
 
 
 
 
 
 
 

1
BIOL 230 Midterm I, 2/12/15 Name_____________________________________
3.  Darwin’s  description  of  natural  selection  characterizes  nature  as  a  ‘wise  force’.  Based  on  our  
discussion  in  recitation,  why  is  this  incorrect?  (5pts)  
 
 
 
 
 
 
 
 
 
 
 
 
4.  In  a  future  lecture  on  adaptation,  we  will  discuss  shell  color  polymorphism  in  the  genus  
Cepaea  (land  snails).    By  1977,  this  system  was  described  as  “a  problem  with  too  many  
solutions”:  there  were  so  many  hypotheses  for  the  maintenance  of  shell  color  variation  that  it  
bordered  on  the  ridiculous.  One  of  these  hypotheses  was  that  predation  by  birds  (thrushes)  
maintains  phenotypic  variation.    Intrigued,  you  set  out  to  test  this  hypothesis  in  a  natural  
population.    At  your  field  site,  you  observe  three  discrete  phenotypes:  yellow,  light  brown,  and  
dark  brown  snails.    From  the  literature  you  find  that  the  yellow  phenotype  is  equivalent  to  a  
homozygous  recessive  (aa),  the  light  brown  phenotype  is  the  heterozygote  (Aa),  and  the  dark  
brown  phenotype  is  the  homozygous  dominant  (AA).    You  painstakingly  collect  a  random  sample  
of  snails  and  place  them  in  an  experimental  enclosure,  such  that  they  cannot  escape  and  are  
exposed  to  bird  predation.    You  allow  bird  predation  to  ensue,  get  commended  by  BIOL230  
students  for  posting  the  gory  videos  on  YouTube,  and  tabulate  the  mortality  events.    You  observe  
that  100  yellow,  200  light  brown,  and  100  dark  brown  snails  were  eaten  by  birds.    Are  your  
results  consistent  with  the  hypothesis  that  bird  predation  acts  as  a  selective  force  on  shell  color  
in  Cepaea?    Why?    Clearly  support  your  answer  qualitatively  and  quantitatively  (10  pts.)  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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BIOL 230 Midterm I, 2/12/15 Name_____________________________________
5.  Consider  a  group  of  many  completely  isolated  subpopulations,  each  of  small  effective  
population  size.    The  initial  allele  frequencies  in  each  subpopulation  are  p  =  q  =  0.5.    After  
10,000  generations  with  no  gene  flow  and  no  selection,  what  will  happen  to  1)  the  mean  allele  
frequency  across  all  subpopulations,  2)  the  variance  in  allele  frequency  across  all  
subpopulations,  3)  the  average  heterozygosity  within  each  subpopulation  as  time  goes  on  
(answers  like  “increase”,  “decrease”,  “stay  the  same”  are  sufficient).    Clearly  state  why.  (10pts)  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
6.    Linnaeus  developed  the  system  of  binomial  nomenclature  that  is  used  in  the  classification  of  
organisms;  a  given  lineage  is  assigned  a  species  name  and  a  genus  name  (e.g.,  Mytilus  (genus)  
edulis  (species)).    This  is  widely  used  in  traditional  classification  schemes,  in  which  genera  are  
organized  into  families,  families  into  orders,  orders  into  classes,  and  classes  into  phyla.    Explain  
how  this  classification  system  could  both  reflect  the  views  of  the  essentialists  (species  have  an  
essential  form  and  are  independently  created;  e.g.,  Cuvier)  and  the  views  of  the  evolutionists  
(lineages  exhibit  descent  with  modification;  e.g.,  Darwin).    (10pts).  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
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BIOL 230 Midterm I, 2/12/15 Name_____________________________________
7.  You  collect  a  series  of  populations  of  the  barnacle,  Semibalanus  balanoides,  from  habitats  in  
Iceland.    This  is  an  arctic,  cold-­‐adapted  species,  meaning  that  it  can  readily  tolerate  freezing  
temperatures  but  is  very  prone  to  death  by  exposure  to  high  temperatures.    In  Iceland,  there  is  a  
very  noticeable  difference  in  temperature  between  the  northern  and  southern  sides  of  the  island;  
water  and  rock  temperatures  are  very  cold  in  the  north,  very  warm  in  the  south,  and  are  
intermediate  on  the  east  and  west  facing  sides  of  the  island.    You  sample  three  populations  from  
the  northern  side  of  the  island,  three  populations  from  the  southern  side  of  the  island,  and  three  
populations  from  the  east  side  of  the  island.    You  then  sequence  the  gene  Arginine  kinase  (Ark)  
from  all  individuals  in  these  populations  and  observe  the  frequencies  given  below.    There  are  
only  two  alleles  at  Ark,  Ark-­‐1  and  Ark-­‐2.    
population f (Ark-1) population f (Ark-1) population f (Ark-1)
north 1 0.03 east 1 0.35 south 1 0.75
north 2 0.09 east 2 0.41 south 2 0.81
north 3 0.07 east 3 0.28 south 3 0.69  
 
 
a)  From  these  data  for  Ark  only,  you  calculate  FSR  as  0.0121,  FRT  as  0.336,  and  FST  as  0.343.    
Provide  two  possible  explanations  for  the  observed  pattern  of  allele  frequency  variation  among  
the  sampled  populations.  (8pts).  
 
 
 
 
 
 
 
 
 
 
 
 
 
c)  You  have  waited  patiently  for  the  cost  of  whole  population  genome  sequencing  to  drop,  and  
now  it  has.  You  sequence  a  random  sample  of  individuals  from  the  replicate  northern  and  
southern  populations,  perform  all  the  associated  analyses,  and  now  get  a  global,  average  Fst  for  
all  variable  positions  in  the  barnacle  genome.  Your  genomic  Fst  estimate  is  0.00001,  and  the  
confidence  intervals  do  not  include  0.343  (in  other  words,  Ark  is  significantly  different  and  not  
an  average  gene).  What  does  this  tell  you  about  the  patterns  of  allele  frequency  variation  at  Ark  
that  you  originally  observed?  (7pts).  
 
 
 
 
 
 
 
 
 

4
BIOL 230 Midterm I, 2/12/15 Name_____________________________________
8.    What  was  the  first  proposed  mechanism  for  evolutionary  change?    Describe  one  way  in  which  
this  mechanism  was  similar  to  Darwin’s  mechanism  of  natural  selection,  and  one  way  in  which  it  
was  distinct.    (10pts.)  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
9.    Briefly outline why the shifting balance model, proposed by S. Wright, relies on a balance between
the evolutionary “forces” of genetic drift, migration/gene flow, and selection. (10pts)
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
5
BIOL 230 Midterm I, 2/12/15 Name_____________________________________
10.    You design a series of experiments to artificially select for high temperature tolerance in
Drosophila melanogaster. In each of the first 10 generations in which selection is applied, you are
able to increase the temperature that the experimental flies can tolerate in a standard thermal
knockdown assay. As a consequence of the artificial selection, what has happened to narrow sense
heritability over the course of the experiment, and why has this occurred? (7 pts)  
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
 
11.    In  natural  populations,  why  does  heritability  for  fitness-­‐related  traits  not  go  to  zero,  despite  
the  continued  action  of  natural  selection?  (8pts).  
 
 
   

6
BIOL 230 Midterm I, 2/12/15 Name_____________________________________
 
 
Formulas   𝑤 = 𝑤!! 𝑝! + 𝑤!" 2𝑝𝑞 + 𝑤!! 𝑞!  
  𝑤!! 𝑝! + 𝑤!" 𝑝𝑞
Population  Genetics   𝑝!!! =  
𝑤!! 𝑝! + 𝑤!" 2𝑝𝑞 + 𝑤!! 𝑞!
  ∆p  =  spq2  /(1  -­‐  sq2)    
P  =  f(AA)  =  p2     𝑝 =  𝑃 + !!𝑄     ∆q  =  -­‐sq2(1  -­‐  q)  
Q  =  f(Aa)  =  2pq   𝑞 =  𝑅 + !!𝑄   p*  =  t/(  s+t)    and    q*  =  s/(  s+t)  
R  =  f(aa)  =  q2    
 
D0  =  p11p22  –  p12p21  
P  +  Q  +  R  =  1     p  +  q  =  1   Dt  =  D0  (1-­‐r)t  
 
   
 
𝐻! − 𝐻!
𝐹! =   Δ𝑞 = 𝜇(1 − 𝑞) − 𝑠𝑞! (1 − 𝑞)  
𝐻! !
! !!
𝐹!" = ! !     𝑞∗ = !
 
!!
 
𝐻! − 𝐻!  
𝐹!" =  
𝐻! Δ𝑞 = −𝑚(𝑞!"  –  𝑞!" ) − 𝑠𝑞!! (1 − 𝑞! )  
𝐻! − 𝐻! 1 𝑝∗ = !!  and    𝑞∗ = !!  
𝐹!" = ≈  
𝐻! 4𝑁! 𝑚 + 1  
 (1  –  FSR)  (1  –  FRT)  =  1  –  FST   s  >  1/(2Ne)  
 

   
𝑝𝑞 Quantitative  Genetics  
𝑉∗ =    
4𝑁! 𝑚 + 1
  P  =  G  +  E  
   
4(𝑁! 𝑁! ) G  =  A  +  D  +  I  
𝑁! =    
𝑁! + 𝑁!
  VP  =  VG  +  VE  
1 1 1
=    
𝑁! 𝑡 𝑁!
  VG  =  VA  +  VD  +  VI  
   
  h2  =  VA/VP  (narrow)  
𝑝! = 𝑝! 1 − 𝜇 ! + 𝑞! (𝜐)!    
𝑞! = 𝑞! 1 − 𝜐 ! + 𝑝! (𝜇)!   h2  =  VG/VP  (broad)  
p*  =  v/(u  +  v)      and      q*  =  u/(u  +  v)    
  R=h2S  
   
𝑝!,!!! = 𝑝! 1 − 𝑚 + 𝑝! (𝑚)  
∆p  =  -­‐m(pxt  –  pyt)  
𝑝!,!!! = 𝑝! 1 − 𝑚 + 𝑝! (𝑛)  
𝑝!,!!! = 𝑝! 1 − 𝑛 + 𝑝! (𝑚)