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Transparency Form of crypsis Involves modification of whole organism Found mostly in pelagic animals Across many taxa. Refractive index (n): Measure of how much the speed of light is reduced in a given medium relative to a reference medium – usually the speed of light in a vacuum (n=1). .
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Transparency Form of crypsis Involves modification of whole organism Found mostly in pelagic animals Across many taxa
Refractive index (n): Measure of how much the speed of light is reduced in a given medium relative to a reference medium – usually the speed of light in a vacuum (n=1). Example: Refractive index of (fresh) water n= 1.333 or 1/1.33 = ¾ the speed of light in a vacuum.
The region below the gray line has a higher index of refraction, and so light traveling through it has a proportionally lower phase speed than in the region above it
Transparent animals / objects • Do not absorb or reflect light
PHYLOGENETIC DISTRIBUTION Pelagic rare w/in group Pelagic common Transparency rare Transparency common
PHYLOGENETIC DISTRIBUTION • Transparency • Appears to have evolved multiple times • Found in most major animal phyla • Primarily pelagic
ECOLOGICAL DISTRIBUTION • Terrestrial– extremely rare • Reflection • Air low refractive index • Difference in ‘n’ between object and surrounding medium • surface reflection decreased transparency • UV • Protective pigmentation • Gravity • Skeletal structures
ECOLOGICAL DISTRIBUTION • Benthic – rare • Match substrate • Pigmentation less costly than transparency? • Shadows Transparent animals may still cast shadows – seen in benthic environments, not in pelagic ones
ECOLOGICAL DISTRIBUTION • Neustonic – rare • Match upwelling light • Blue or brown • works from above but not from below • UV • Protective pigmentation
ECOLOGICAL DISTRIBUTION • Aphotic – rare • Red or Black pigmentation • Absorb bioluminescent light
ECOLOGICAL DISTRIBUTION MOST TRANSPARENT ANIMALS 10 MAJOR GROUPS (Pelagic): Cubozoans Hydrozoans Ctenophores (non-beroid) Hyperiid Amphipods TomopteridPolychaetes Heteropods Pteropods Cranchiid squid Thaliaceans Chaetognaths
INTERACTIONS • Transparency of an animal depends on • Fraction of light that passes through • Not absorbed or reflected (scattered) • Contrast • Brightness of object relative to its background • Decreases with distance • Visual capacity of viewer • Sighting distance • Adaptations to break transparency
INTERACTIONS Adaptations to break transparency: UV vision Polarization vision Viewing angle (behavioral)
INTERACTIONS Adaptations to break transparency: UV vision Increased scattering of light in UV range increased contrast
INTERACTIONS Adaptations to break transparency: Polarization vision Can detect changes in polarization of highly polarized oceanic light
INTERACTIONS Adaptations to break transparency: Viewing angle (behavioral) Snell’s Window condensed horizon effect increases contrast of transparent objects outside of “window”
ADAPTATIONS for TRANSPARENCY • Most organic molecules do not absorb light • Transparency is a matter of reducing light reflections or scattering caused by light passing through media with different refractive indices.
ADAPTATIONS for TRANSPARENCY Transparent animals must compensate for their varied constituent refractive indices
ADAPTATIONS for TRANSPARENCY • Macro • Cloaking of non-transparent features • Eyes • Compact retinas • Mirrors • Counterillumination • Separation of eyes • Guts • Elongated • Vertically oriented (decreases view from above/below) • Mirrored/reflective • Counterilluminating bioluminescence – minimizes shadows
ADAPTATIONS for TRANSPARENCY • Macro • Cloaking of non-transparent features • Eyes • Guts • Be flat • Light attenuation decreases exponentially as tissue thickness decreases (Thinner = more light passes through)
ADAPTATIONS for TRANSPARENCY • Micro • Surface • Extracellular Matrix • Cellular
ADAPTATIONS for TRANSPARENCY • Micro • Surface • Moth eye surfaces • Bumps have widths <1/2 the wavelength of incident light Create a refractive index gradient Decreases effective surface refractive index Decreases scatter
WIDTH OF BUMP IS LESS THAN HALF A WAVELENGTH OF LIGHT INDEX OF REFRACTION DARK = BUMPS LIGHT = SURROUNDING MEDIUM
ADAPTATIONS for TRANSPARENCY • Micro • Extracellular Tissues • Average refractive index constant over distance ½ the wavelength of incident light • Low scattering • Due to destructive interference of scattered light
ADAPTATIONS for TRANSPARENCY • Micro • Extracellular Tissues • Average refractive index constant over distance ½ the wavelength of incident light • Low scattering • Due to destructive interference of scattered light Caused by densely packed similar objects Example: Mammalian cornea and lens tissues densely packed so scatter is ordered and reduced
Transparency and biomechanical properties of the cornea depend on the structure and organization of corneal stroma. Collagen fibers and fibers interconnecting to the network formed collagen bundles, which were regular and parallel to the corneal surface Barbaro, Mol Vis 2009; 15:2084-2093. http://www.molvis.org/molvis/v15/a224
ADAPTATIONS for TRANSPARENCY • Micro • Cellular Tissues • More complex • Necessary components with different refractive indices
ADAPTATIONS for TRANSPARENCY • Micro • Cellular Tissues • More complex • Necessary components with different refractive indices • Theoretical model: • Size matters • Distribution • Refractive index • Shape does not matter that much
ADAPTATIONS for TRANSPARENCY • Micro • Cellular Tissues • Theoretical model: • Size matters • Distribution • Refractive index • Shape does not matter that much • Theoretical predictions: