2. LOW LIGHT, BAD FLIGHT
VISION AND VISUAL NAVIGATION IN
NOCTURNAL INSECTS
Prashant
PAL 0013
2
3. Compound eyes:
Insects recognize and react to conspecifics
Distinguish and avoid predators
Locate food sources and intercept prey
Navigate
Walk, swim or fly
Nocturnal insects:
Introduction
Light levels can be up to 11 orders of magnitude lower
3
8. Visual processing in nocturnal
insects
Vision
Eyes with an enhanced optical sensitivity to light
Visual neurons that sacrifice spatial and temporal resolution
to improve visual reliability for the slower and coarser
features of the world
EYES AND VISION IN NOCTURNAL INSECTS
Optics
Retina of the compound eye
8
9. A superposition eye can have
optical sensitivity 100–1000 times
higher than that of an apposition
compound eye
The Optical Designs of Nocturnal Compound Eyes
A
9
10. The ratio of the number of photons absorbed by a photoreceptor
to the number emitted per steradian of solid angle from a unit
area of an extended source
Where,
A-Diameter of eye operture
l - length of the rhabdom
K -peak absorption coefficient
of the visual pigment
f -focal length of the
ommatidium
d -diameter of the rhabdom.
Optical sensitivity
10
11. Good sensitivity to a spatially extended scene results from
a pupil of large area (π A2/4)
Photoreceptors that each view a large solid angle of visual
space (πd2/4 f 2 steradians)
Absorb a substantial fraction of the incident light (kl/2.3+kl).
This equation predicts
11
12. M. genalisA. mellifera
2 μm 8 μmRhabdum width
Facet size 36 μm20 μm
M. genalis an optical sensitivity that is roughly 27 times
greater than that of A. mellifera
Warrant et al., 2004
How nocturnal life has affected the optical structure
and sensitivity?
12
13. Warrant et al., 2006
Ocellar optics in nocturnal and diurnal bees and wasps
Nocturnal sweat bee, Megalopta genalis,
Nocturnal paper wasp, Apoica pallens
Diurnal paper wasp, Polistes occidentalis
13
18. General property of photoreceptors- Bumps
At higher intensities: the bump responses fuse to create a
graded response whose duration and amplitude are
proportional to the duration and amplitude of the light
stimulus
Photoreception and the Reliability of
Vision in Dim Light
18
19. At very low light levels: a light stimulus of constant intensity is
coded as a train of bumps
At somewhat higher light levels: the constant intensity is coded
by a graded potential of particular amplitude 19
20. The major limitation for nocturnal vision in insects
Arises from the stochastic nature of photon arrival and
absorption
Sources of Visual noise
Photon shot noise
Dark noise
Transducer noise
Visual noise
20
21. • Photoreceptor absorbing a number of N photons
experiences an uncertainty (or photon shot noise) of √N
photons
(Land, 1981; Warrant and McIntyre, 1993)
• Decreasing photon catch in dim light results in an
increasing noise level
• As two visual channels need to detect sufficient photons in
order to reduce this noise level
PHOTON SHOT NOISE
21
22. Consists of spontaneous thermal responses in the
absence of photons, which are indistinguishable from
membrane potentials (quantum bumps) produced by photons
(Barlow, 1956)
These fluctuations are more frequent at higher
temperatures and introduce uncertainty at low light
intensities.
Even though dark noise is much lower in invertebrates than
in vertebrates
Dark noise
22
23. Photoreceptors are incapable of producing identical bumps
of fixed amplitude, latency and duration to each (identical)
photon of absorbed light.
This source of noise, originating in the biochemical
processes leading to signal amplification
To maximise the photon catch or signal-to-noise ratio to
enhance sensitivity
Transducer noise
23
24. Photoreceptor responses to single photons (i.e bumps)
are much larger in nocturnal insects
Retinal adaptations for nocturnal
vision
Large bumps have been demonstrated
Nocturnal crane flies
Cockroaches
Bees
Spiders
24
25. Quantum bumps of nocturnal M. genalis and diurnal L.
leucozonium 25
26. Contrast gain of the bees M. genalis and L. leucozonium
Fredriksen et al., 200826
27. A large number of animals are known to use colour to
detect, discriminate and recognise objects
Food sources
Mating partners
Landmarks or their homes
Nocturnal Color Vision
An animal needs to possess and use at least two types of photoreceptors,
with different spectral sensitivities, to look at an object
27
28. Simple 4-stage model of colour discrimination with two
spectral types of receptors.
28
30. UV
Violet
Green
Kelber, 2003
Deilephila elpenorMacroglossum stellatarum
Colour Vision in Diurnal and
Nocturnal Hawkmoths
Three different spectral classes of photoreceptors
30
31. Schematic drawings of the structure of the rhabdom of
Deilephila elpenor
Schlecht et al., 1978
31
32. Color vision in D. elpenor is color-constant
This moth can not only be trained to associate a sugar
reward with a blue disc at starlight
Discriminate this blue disk from other discs in various
shades of gray with a choice frequency of at least 80%.
Kelber, 2003
32
35. Many nocturnal insects have evolved sufficiently
sensitive visual systems
Celestial cues
Terrestrial visual Cues
Nocturnal Navigation and
Orientation
35
36. At night, the brightest and most easily discernable cue in
the sky is undoubtedly the moon
Navigation and orientation using
celestial cues
Its bright disk is used for orientation and navigation in a number of
different nocturnal insects
Ants Earwigs
Moths
Beetles
36
37. A much dimmer and more subtle cue associated with the
moon is its pattern of polarized light.
This circular pattern, centered around the moon, arises
because of the atmospheric scattering of moonlight as it
travels toward Earth
Light is most polarized around a circular celestial locus
90◦ from the moon, and the circular pattern of polarized
light moves with the moon
Cont….
37
38. On full moon nights
On four nights before and after this event
Dacke et al., 2003
Lunar orientation in a beetle
38
39. The path taken by a ball-rolling Scarabaeus zambesianus
0 to 90 90 or 180
39
40. Circular diagrams of turns made by Scarabaeus zambesianus
rolling under the night sky.
40
41. Visual detection of optic flow is also clearly necessary for
controlling nocturnal flight
Navigation and orientation using
terrestrial cues
Dim light gypsy moths
Mosquitoes
Locust
41
43. X. leucothorax is diurnal
X. tenuiscapa is largely diurnal and
occasionally crepuscular
X. tranquebarica is truly nocturnal
Somanathan et al., 2008
Flight activity in three species of carpenter bees
in relation to light intensities
43
44. Somanathan et al., 2008
Flight activity in all three species as a function of light
intensity
44
46. X. leucothorax X. tenuiscapa
X. tranquebarica
46
Scanning electron micrograph of heads
47. Canopy or Individual trees
As the animal moves under the tree canopy, the brighter
sky in the gaps of the canopy, together with the darker
area under the canopy
Ex: Nocturnal shield bug, Parastrachia japonensis
Insects living in forests
47
54. A) initial orientation of individual ants at the nest the
B) time taken to exit the 30cm circle and
C) the proportion that crossed the 1.2m reference line 54
56. Neural adaptations
An increase in the response gain of the
photoreceptors with decreasing light intensity can
further enhance sensitivity but does not improve
photon capture itself
(Laughlin, 1981)
The ultimate solution to optimise sensitivity at
low light intensities is to process the incoming
visual signal using a strategy of neural summation
in space and time
56
57. Temporal Summation
Visual systems can also improve image reliability
at night by slowing vision down
Lengthening the eye’s visual integration time at
night
Signal-to-noise ratio of lower temporal
frequencies is improved
57
58. Temporal summation results in a slower but more reliable
visual world
Extremely long photoreceptor integration times
Sit-and-wait predators and slowly moving animals,
temporal summation is certainly a good strategy
Cont…
58
60. Photons are integrated over wider visual fields,
which is similar to a widening of the angular
sensitivity function
Only when neural summation is matched to the
extent of the visual overlap present in the eye
60
62. Greiner et al., 2004 and Ribi, 1975
Comparison of the First-Order Interneurons, L-fiber types L3
and L4, of the M. genalis female (left) and the worker
honeybee A. mellifera (right)
62
63. Nocturnal insects have excellent night vision
With the capacity to discriminate colors
Orient themselves using faint celestial cues fly unimpeded
through a complicated habitat
Navigate to and from a nest using learned visual
landmarks
Conclusion
63
64. The photoreceptors of nocturnal insects respond more
slowly and have a higher contrast gain
A neural strategy of spatial and temporal summation at a
higher level in the visual system is hypothesized as the
necessary bridge between retinal signaling and
visual behavior.
Cont…
64
The large rhabdoms of Megalopta are clearly an adaptation for improved sensitivity at night, and the interesting observation that the equally nocturnal Apoica has small rhabdoms suggests that some of their sensitivity may have been offered in favour of spatial resolution. This conclusion is reinforced by calculations of the occupation ratio of rhabdoms in the retina (the percentage cross-sectional area of rhabdoms within a unit cross-sectional area of retina). The occupation ratios of the wasps Apoica and Polistes are quite small (9.5% and 6.1%, respectively: Table 1). Even though Apoica is nocturnal, its rhabdom occupation ratio is only slightly larger than that of its diurnal relative. In contrast, the nocturnal bee Megalopta has a rhabdom occupation ratio over three times greater (33.6%: Table 1).
At the level of the photoreceptors, the reliability of vision is determined by the repeatability of this response
They were then displaced either 2m to the left or 2m to the right of the nest-to-tree line (R). Normal foraging paths are shown in grey, displaced paths are in black.