eye evolution (guest lecture ) - claremont collegesfaculty.jsd.claremont.edu/dmcfarlane... ·...
TRANSCRIPT
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Eye evolution (guest lecture )
Lars Schmitz 10/25/2012
Why don’t plants have eyes?
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
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… about a third have light-sensitive
cells.
… about a third have proper eyes.
… about a third have no visual
specializations whatsoever.
Of the ~30 animal phyla …
“proper” eye: an organ able to compare light
intensities in different directions [spatial vision]
Turbellarian Flatworm
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Chambered Eyes
Scallop
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Vertebrate eyes
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Amniote Eye
California Condor
cornea
lens
iris
pupil
Retina
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Chambered Eyes
everse (octopus)
vs.
inverse (cod)
Cephalopod eyes
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Compound Eyes
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schizochroal
holochroal
abathochroal
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
To suppose that the eye, with all its inimitable contrivances for adjusting the focus to different distances, for admitting different amounts of light, and for the correction of spherical and chromatic aberration, could have been formed by natural selection, seems, I freely confess, absurd in the highest possible degree. (Darwin 1872)
Darwin on eye evolution
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Yet reason tells me, that if numerous gradations from a perfect and complex eye to one very imperfect and simple, each grade being useful to its possessor, can be shown to exist; if further, the eye does vary ever so slightly, and the variations be inherited, which is certainly the case; and if any variation or modification in the organ be ever useful to an animal under changing conditions of life, then the difficulty of believing that a perfect and complex eye could be formed by natural selection, though insuperable by our imagination, can hardly be considered real. (Darwin 1872, 143-144)
Darwin on eye evolution (next paragraph)
Time required to evolve an eye
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
Refractive
System
Receptors Neural Processing
(Retina + Brain)
Behavioral
Anatomical
Optical
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Visual
Capacity Resolution
Field of View
Sensitivity
Light PND
S
Quantifiable as
Spatial Sampling Frequency (SF)
PND: Posterior nodal
distance
S: Separation of
receptor centers
(min. ~2 µm)
SF = ____ PND
2·s
Land (1981)
Resolution
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Land and
Nilsson 2002 Spatial Sampling Frequency
Octopus
Aeschna
Nautilus
Aquila audax
Falco berigora
resolving power
behavioral: 132-143 c/deg
anatomical: 140 c/deg
resolving power
behavioral: 69-76 c/deg
anatomical: 76 c/deg
Reymond 1985,1987
(Wedge-tailed Eagle)
(Brown Falcon)
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Accommodation
Visual
Capacity Resolution
Field of View
Sensitivity
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Sensitivity to Point Light Sources
klr eer
EAN 1
16 2
2Sensitivity ~
monochromatic light
E: # of photons originating at source l: length of receptors
A: aperture k: peak absorption coefficient r: distance to light source α: combined attenuation coefficient
for given receptors, light distance and intensity
A2
Warrant (1999)
kl
klA
PND
d3.2
2
2
2
Sensitivity=
PND
A2
d: diameter of receptors
l: length of receptors
k: peak absorption coefficient
A: aperture
PND: posterior nodal distance
(for given receptor dimensions)
Land (1981)
Sensitivity to Extended Sources
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f-number
PND
A2
Sensitivity ~
f-number = PND
A
the lower the better
PND PND
A S
TRADE OFF
Resolution Sensitivity
SF = ____
2·s
PND F =
____ PND
A
for given eyeball size,
receptors, neural processing
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Visual
Capacity Resolution
Field of View
Sensitivity
Visual Fields
Martin et al. 2007
Apteryx sp.
(Kiwi)
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Steatornis caripensis
(Oilbird)
38°
Visual
Capacity Resolution
Field of View
Sensitivity
Contrast sensitivity
improved by 40% Visual threshold
improved by 40%
Trade Off
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
Bio-mechanics
Bio-optics
EVOLUTION
Physical constraints
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Light Levels Luminance [photons m-2 sr-1 s-1]
8 lo
g u
nit
s
water depth [m]
terrestrial marine 1
4 lo
g u
nit
s
Po
int
Lig
ht
So
urc
es
4
Starlight
Moonlight
Bright
Sunlight
6
10
8
12
14
16
18
20
1000
800
600
400
200
0
Epipelagic
Mesopelagic
Bathypelagic
Oil Bird
Scopelarchus guentheri
Martin et al. 2004
Warrant 2004
Retinal adaptations
Tapetum Lucidum IRS
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Tapetum Lucidum
Retinal adaptations
Apteryx mantelli
kiwifoundation.org.nz
inside
outside
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Retina
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Muraenidae
perco-
morphs
holo-
centrids elopo-
morphs
Holocentridae
perco-
morphs
holo-
centrids elopo-
morphs
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Apogonidae
perco-
morphs
holo-
centrids elopo-
morphs
Priacanthidae
perco-
morphs
holo-
centrids elopo-
morphs
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Sciaenidae
perco-
morphs
holo-
centrids elopo-
morphs
Pempheridae
perco-
morphs
holo-
centrids elopo-
morphs
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Haemulidae perco-
morphs
holo-
centrids elopo-
morphs
diurnal nocturnal
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Schmitz and Wainwright 2011 (BMC Evol. Biol.)
A
D x f
2
small large
A: Aperture
D: retinal area
f: focal length
aquatic
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Schmitz and Wainwright, 2011 (BMC Evol. Biol.)
Eye morphospace of reef fish
round, large
pupil;
large lens
large, elliptical
pupil;
small lens
small, elliptical
pupil
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Light sensitivity
large, elliptical
pupil;
small lens
round, large
pupil;
large lens
small, elliptical
pupil
Diurnal
IRS
IRS
IRS
Collared Lizard
Chipmunk
King Parrot
Flame Robin
Eastern Robin
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Leopard Gecko
Nocturnal
Crested Owl
ADW
IRS
Oilbird stanford.edu/~cagan/southamerica.htm
Brown Rat ADW
Cathemeral Crepuscular
Asian Elephant
BNAO
Common Nighthawk
BNAO
Giraffe giraffe.org
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A
D x f
2
small large
A: Aperture
D: retinal area
f: focal length
terrestrial
Eye morphospace
Discriminant Axis 1
Dis
cri
min
an
t Axis
2
95% confidence ellipses
mammals, n=35
birds, n=15
squamates, n=8
Variables: eyeball diameter,
axial length,
lens diameter
Schmitz and Motani 2010, Vision Res.
misclassified
proportion (pFDA):
13.4%
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
Weishampel et al. 2004
-mixed time periods-
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Weishampel et al. 2004
-mixed time periods-
Scleral Rings
Bubo
Osteology
and optics?
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Mammal Bird
Scleral ring
Scaphognathus
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Lens diameter
External scleral ring
Internal scleral ring
Axial length
Eyeball diameter Orbit length
Hard-tissue Soft-tissue
Orbit length External + internal
scleral ring
diameter
Schmitz 2009 (J. Morphol.)
diurnal
nocturnal
Schmitz and Motani 2011 (Science)
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Dinosauria
Archosauria
33 Mesozoic archosaur species
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Weishampel et al. 2004
-mixed time periods-
cathemeral/
crepuscular
equivocal (either
nocturnal or cathemeral/
crepuscular)
diurnal
nocturnal
8
14
9
2
Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
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Point light sources
Extended luminous sources (form illumination)
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Extended non-luminous sources
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Questions…
• How many times did eyes evolve?
• How long did it take to evolve an
eye?
• What makes a good eye?
• How does ecology influence the
evolution of eyes?
• Were some dinosaurs nocturnal?
• Why do giant squid and ichthyosaurs
have giant eyes?
• What fish have reduced eyes?
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Scytalina
Forbesichthys
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Ditropichthys
Melanocetus
Oneirodes
Bufoceratias