Ecology Biology 3500
                                                                                                                Dr. Adams

Chapter 4: Population Genetics and Natural Selection
    You must know definitions of the following:  evolution, natural selection, adaptation,
        genes/alleles, and genetic terminology (dominant/recessive, homozygous/heterozygous,
        incomplete dominance, loci, etc.)

    Evolution by Natural Selection -- (VALID) Assumptions:  (includes time)
        1.  Reproduction with variation.
        2.  At least some of the variation is heritable
        3.  Overproduction of offspring (leads to competition, more predation/disease, death)
        4.  The variation means that some will have a higher chance of survival than others.
   Leads to adaptations allowing "survival of the fittest" --  what IS fitness?

Variation within populations
    Plants -- Cinquefoil examples
        Potentilla glandulosa -- significant variation across altitudinal gradient.
            If completely environmental, variation would disappear in common garden experiments.
            When grown in common gardens, not all variation disappears, and those adapted to a
                particular altitude largely grew best at their own altitude, though the middle elevation
                individuals appeared to excel at all altitudes -- clearly some selected genetic differences
                that enhanced survivorship within certain habitats - these are called ecotypes.
        Potentilla nivea and pulchella complexes (on Spitsbergen Island, Norway)
            P. pulchella shows significant differences (three morphs) in different habitats; common
                garden studies, however, indicate that the differences are due to a plastic genome
            The P. nivea complex, thought to include three species, indeed does when genetic
                analysis taken into account.
        So, in one genus there is a species (pulchella) that shows variation that is virtually completely
            environmental, another species (glandulosa) which has some genetic basis for the
            morphological differences between populations, and a species complex (nivea) where
            the differences are significant enough that there are three distinct species
    Animals -- Whitefish (Coregonus sp.) in isolated rivers and lakes in the Alps
        Phenotypic and genetic analysis of specimens from 19 described populations indicate that
            using the data collectively gives a roughly 80% ability to assign individuals to appropriate
            source populations -- in other words, there is some significant genetic distinctness.  Enough
            to be called species?  The investigators went so far as to call them "evolutionarily
            significant units," enough so that they should be managed separately (not mixed).

Hardy-Weinberg -- for a trait with two alleles within a population:
    Equations:     p = frequency of A       q = frequency of a    p + q = 1 (obviously)
                p2 = frequency of AA      2pq = frequency of Aa     q2 = frequency of aa
                    Again, clearly p2 + 2pq + q2 = 1
        (I will show you the derivation of this equation in class, even though you probably know
            how it is derived already; we will also do an example or two or . . . ?)

    The idea here is that a population would be considered to be in H-W equilibrium if the
        equations resulted in actual representation of a real population, and the frequencies didn't
        change from generation to generation.  To be in H-W equilibrium, however, the population
        would have to exhibit the following:
      1.  Random mating
      2.  No mutation
      3.  Large population size -- prevents chance events (genetic drift) from altering frequencies
      4.  No immigration or emigration -- in other words, populations are in isolation, and have NO
            gene flow.
      5.  No selection -- in other words, all organisms have equal fitness

        How many of these conditions are met in actual populations?  VERY few are met, and most
certainly not all, in ANY population.  As such, genetic frequencies will change, which means . . .
Natural Selection (H-W requirement #5): understand that selection is not necessarily happening 
    continuously in one direction -- what is favorable now may not be favorable later; it is not a
    process of perfection; it may act on different populations of the same organism differently
    (different selective pressures)
  Types of selection:
    1.  Stabilizing -- extremes selected against
    2.  Directional -- entire curve shifts to one side or other; probably most common
    3.  Disruptive -- both extremes selected for; typically doesn't happen within one population
            (though it can); this usually happens in different populations of a species, which can
            lead to divergence and new species

    Heritability -- The fraction of the variation is due to variation in genes, represented by

h2  =  Heritability  =  VG   =      VG__       P  = phenotypic, G = genetic, E = environmental
                                           VP       VG + VE

If heritability is near 0, then that means . . . ?  If heritability is .50, then that means . . . ? 
   If heritability is near 1, then that means . . . ?

        Understand that natural selection working on traits with low heritability will NOT result in
any significant genetic change, certainly not in the short term. 
        Which brings us to an important question.  Can adaptation take place quickly in a trait with
significant heritability?  This, of course, is the meat of the idea of evolution.

  Stabilizing Selection:  Egg Size in Ural owls (and other birds); human birth weight
  Directional Selection:   Beak lengths of Soapberry Bugs and introduced foodplants; you will be
            responsible for the story in both the U.S. and Australia.
        Turns out that heritability is high -- juveniles reared on one hostplant retained beak length
            when switched to another.  Natural selection has adapted different populations of these
            bugs within 30 to 100 years.
  Disruptive Selection:  Darwin's Finches on the Galapagos, butterfly species

Change Due to Chance
    Genetic Drift -- more potent in smaller populations (as you will see in lab)
        (remember that mutation is, in essence, another chance event)
    Genetic Variation in island populations -- almost always less than variation in mainland pops.
    Remember that "islands" can also apply to (semi-) isolated populations in any kind of habitat.
        In reduced patches of habitat with small populations, inbreeding also reduces variation.
            Example:  Glanville Fritillary Butterfly in Finland (see pages 94-95)

Chapter 5:  Temperature Relations
    Local variation in temperatures due to: altitude, aspect, vegetation,  ground color, 
        topographic relief (boulders/burrows), nearby water (riparian habitats and vegetation)
            In aquatic environments, depth in the water minimizes temperature fluctuations, and,
                as indicated previously, water in general buffers changes in temp.

    An evolutionary trade-off:  Adapting to one set of circumstances typically minimizes the
        ability of organisms to succeed in other environments. They must choose to allocate
        resources so that they can maximize their performance in their chosen environment.

    Organismal performance -- most organisms adapted to a rather narrow range of conditions
        for their activities, including a rather narrow range of temperatures (though homeothermy
        provides much greater temperature tolerance by providing a narrow internal temperature
        range).  Organisms can allocate only so much for each activity, and therefore less
        temperature stress leaves more energy for other activities. Why is a narrow temperature
        range useful??  Enzyme function. 
            Questions to ponder:  1.Can different organisms have different enzymes to do the
                same process but function at different temperatures? and 2. Can the SAME org.
                have more than one enzyme to do the same process but function at dif. temps.?
                3.  Can the same organism acclimate to different environmental conditions?
        Photosynthetic efficiency peaks in plants from dif. latitudes/altitudes at dif. temps.
            This trend is repeated for virtually any group of organisms.
        Endothermy increases range, but requires more energy input (see below)

    Regulating temperature -- an attempt to balance heat gain vs. heat loss
            Sources of heat/heat loss: metabolism (g), conduction (g or l), convection (g or l),  
(g or l), evaporation (transpiration for plants) (l)
                Poikilothermy (varies with ambient), Homeotherms (constant TB)
                Ectothermy and Endothermy
        Plants: Different strategies for different habitats --
            Deserts: little transpiration (why?); leaves narrowed/reflective/off the ground (why?)
            Arctic/Alpine: opposite of deserts in many respects:
                    Leaves flattened/darkened/near ground; can reach temps far above ambient
            Tropical Alpine plants and giant rosettes
            Thermogenic plants (skunk cabbage; see page 119)
            Many ectotherms (eg., lizards, beetles) bask in cool environs, stand "tall" and 
                "dance" in hot locations. Insects tend to be darker in cool climes, lighter in warm
                (dif. broods may vary with seasons and with dif. tempse in the environment in which
                they grow; see grasshoppers in Fig. 5.20).
            Endotherms do have dif. (but higher than ectotherms) metabolic rates depending on
                preferred habitat.  Aquatic endotherms typically have significant insulation.
                Interestingly, insects (and others) can act as endotherms with muscular thermo-
                genesis (shivering).    Know the concept of thermoneutral zones.
            Endothermic aquatic animals (some fish [tuna/sharks], mammals, penguins)
                Mammals and penguins -- no large respiratory surface exposed to the water; thick
                       feathers/fur/blubber protect them from heat loss
                Fish endotherms can swim faster and greater distances -- more access to prey
                      Apparently can maintain temp with heat produced by highly active muscles

             Interestingly, some insects can use muscular thermogenesis to raise body temps
                     significantly above the surrounding air temps (sometimes as much as 50 deg C)

    Surviving the extremes: inactivity -- torpor, diapause, hibernation/estivation
            Special adaptation in invertebrates -- antifreeze.

Chapter 6:  Water Relations -- water moves down concentration/pressure gradients
    Life is a never ending attempt at balancing water loss with water gain
        For terrestrial organisms, especially in arid environments, it can be the #1 factor
        determining existence in a particular biome

    Water availability
        Atmospheric water -- relative humidity/vapor pressure
            100% humidity = precipitation
            <100% = vapor pressure deficit; when low, water leaves organisms into the air
        Aquatic environments:  You should understand osmosis, osmotic pressure, and
            hypo-/hyperosmotic conditions
.  Will briefly discuss invertebrates and bony fish
            in fresh/marine environments, and cartilaginous fish in marine environment.*
        From soil to plants -- follows a water potential gradient; from soil through xylem to
            leaves and out (transpiration) through stomates/lenticels (a continuous water
            column).  Though stomates are for CO2/O2 exchange, stomates WILL be closed
            to prevent excessive water loss -- so water is the ultimate controller of stomates.

    Water regulation in animals/plants on land
        Again, water losses (how?) must be balanced by water gains (how?)
                      secretions (l), evaporation/transpiration (l), absorption (g)
            Examples (see book, pgs. 133-134)
            Modifications for acquisition/conservation under certain conditions:
                Plants:  water gain from soil (or water); water loss by transpiration and secretions
                    1. more root growth in plants when water stressed, moreso in species found
                            in drier climates
                    2. heavier cuticle on/narrowing of leaves in drier climates
                    3. C4/CAM photosynthesis and stomate narrowing in drier climates
                    4. broad, shallow root in drier climates to acquire water when available (cactus)
                Animals:  water gain by eating/drinking/metabolism; water loss by urination/feces/
                        evaporation (sweating in some)
                    1.  more armor (turtles) in drier climates
                    2.  similarly, thicker, more waterproofed cuticle (tiger beetles) in drier climates
                    3.  activity at night (many mammals, scorpions, etc.); some so efficient at water
                            conservation (Merriam's Kangaroo Rats) that they do not need to drink in
                            desert habitats --subsist entirely on food and metabolic water.
                    4. drink large amounts (camel) when water available in dry climates
                    5. evaporative cooling, even possible in some small arthropods

            Camels and Saguaro cactus
            Scorpions and Cicadas -- WHY does the specific *Cicada species discussed "want" to
                    be active during the daytime in the desert?

    Water and salt balance in Aquatic environments
        *As indicated above, you will need to know what is going on with:   
            Marine Fish (bony and cartilaginous) and marine invertebrates
                Invertebrates largely isosmotic -- no energy expended to maintain body water,
                    though may need to expend some energy to balance certain solutes
                Cartilaginous -- slightly hyperosmotic; gain water through osmosis (across gut, gill
                    membranes), eliminate excesses through (dilute) urine; sodium, too, diffuses in, 
                    with excesses eliminated through a rectal salt gland.
                Bony fish -- hypoosmotic; gain water by constantly drinking, but must rid body of
                    salt picked up with water -- do so with specialized salt glands associated with
                    gills, and by excretion of (concentrated) urine
            Fresh water bony fish and invertebrates -- hyperosmotic
                Bony fish -- like cartilaginous fish in marine environment; easily gain excess water
                    from (and lose salts by diffusion to) external environment.  Get rid of excess water
                    through large amounts of dilute urine; have cells in gills that actively pick up salts.
                Invertebrates -- tissues are between one-half and one tenth as concentrated as
                    marine relatives -- limits of dilution are determined by the minimal levels of solutes
                    in body fluids that must exist for normal nerve, muscle, etc. function.  Like for the
                    fresh water bony fish, must actively pump out water and actively pump in salts.