VISION AND VISUAL NAVIGATION IN
NOCTURNAL INSECTS
PRESENTED BY
T.YUVASRI
23PGA205
CHAIRMAN : Dr. M. KANDIBANE
PANDIT JAWAHARLAL NEHRU COLLEGE OF AGRICULTURE
AND RESEARCH INSTITUTE, KARAIKAL- 609603
INTRODUCTION
❖ Insects play tremendous - ecological and economic role
globally
❖ Insect activity is on average 31.4% higher at night than in
the day
(Wong and Didham, 2024)
❖ Insects have extremely complex vision systems - small
eyes and relatively tiny brains
(Warrant, 2017)
INSECT EYES
❖ Insect eyes are a marvel of evolutionary adaptation
❖ Unlike the simple, single lens eyes - vertebrates, Insects possess compound
eyes
❖ Insect eyes are adapted to various ecological niches-specialized visual systems to
suit their specific needs
(Buschbeck and Friedrich, 2008)
INSECT EYES
EVOLUTION
❖The origin of compound eyes can be
traced back to the Cambrian period - 500
million years ago
❖Modern insect ancestor eyes only detect
light and dark
❖Simple eyespots to compound eyes
involved incremental changes that provided
- Better predator detection and improved
navigation
(Warrant and Nilsson, 2006)
DORSAL
OCELLI
T YPES OF
INSECT EYES
STRUCTURE OF INSECT EYES
LATERAL OCELLI
DORSAL OCELLI
COMPOUND EYE
SIMPLE EYE TYPES
SIMPLE EYES
❖ It comprise of single lens for collecting and focusing light
SIMPLE EYES
LATERAL OCELLI DORSAL OCELLI
LATERAL OCELLI
❖ Larval vision relies on single-lensed eyes called
stemmata
❖ Larval hymenopterans (excluding symphytans) and fleas
do not possess stemmata
(Simpson and Douglas, 2013)
❖ Sawflies and Tiger beetles have more advanced
stemmata with improved visual abilities
(Paulus, 1986)
STEMMATA OF FIREFLY
❖ Photuris larvae - single stemma with each eye containing 88 photoreceptors
❖ Fusion stemmata - Two clusters of cells fuse to form the dorsal stemma, and three ventral
clusters merge to form the ventral stemma
❖ Wide, net-like rhabdomere - vertical columns shared central region - each rhabdom has a
‘fused’ organization - consistent with rhabdoms -maximal photon capture
(Murphy and Moiseff, 2019)
DORSAL OCELLI
✓ Dorsal ocelli - front surface of the head of nymphs and adults of several insect
✓ Present in Bees, locusts and dragonflies
✓ It cannot form an image and is unable to recognize the object
✓ Sensors of ambient light intensity e.g., for the control of circadian rhythms
(Honkanen et al., 2018)
COMPOUND EYE
❖ Compound eyes are the primary visual
receptors of adult insects and larval
hemimetabola
❖ They are constructed from many similar
units called ommatidia
Lepismatidae - 12 ommatidia on each side
Collembola - 8 widely spaced ommatidia
Dragonfly - 30,000
Drone bees - 10,000
Ponera punctatissima - 1
(Simpson and Douglas, 2013)
STRUCTURE OF COMPOUND EYE
OMMATIDIA
CORNEAL LENS
CRYSTALLINE CONE
PRIMARY PIGMENT CELLS
SECONDARY PIGMENT CELLS
RHABDOM
PHOTORECEPTOR
BASEMENT MEMBRANE
PARTS OF COMPOUND EYE
OMMATIDIA
Light-gathering part and a
sensory part, perceiving the
radiation and transforming it
into electrical energy
CORNEAL LENS
❖ The cuticle covering the eye is transparent
and colorless - covers outer ommatidium
❖ It is produced by corneagen cells
❖ Later withdraw to the sides primary pigment
cells
(Zhao et al., 2014)
CRYSTALINE CONE
❖ Crystalline cone and the lens work
together to refract light onto the
rhabdom
(Tomlinson, 2012)
PARTS OF COMPOUND EYE
PIGMENT CELLS
✓ Diurnal insects - pigment
cells surrounding each
ommatidium – Mosaic image
✓ Nocturnal and crepuscular
insects have pigment cells that
do not completely isolate each
facet less distinct mosaic
PRIMARY PIGMENT CELLS
❖ Primary pigment cells in insect
eyes create an insulating
cylinder around the lens unit,
which helps improve visual
acuity
❖ Crescent-shaped
SECONDARY PIGMENT CELLS
❖ Found in the retina of insect
compound eyes - between two
adjacent ommatidia
❖ Regulate the amount of light
that reaches the photoreceptors
(Tomlinson, 2012)
PARTS OF COMPOUND EYE
RHABDOM
✓ Light−sensitive part
✓ Rod-like structure, secreted by retinula cells
and centered on the optical axis
✓ It contains light-sensitive pigments – rhodopsin
✓ Pigments absorbs wavelength of incident
light and generate nerve impulses -
photochemical process
✓ Resolve wavelength and plane of polarization
Fused rhabdom – Bees
Open rhabdom - Fruitflies
(Zelhof et al., 2006)
PARTS OF COMPOUND EYE
PHOTORECEPTOR
✓ Ommatidium - eight to nine photoreceptors
✓ Photoreceptors have rhabdomeres, microvillar
structures where visual pigments are densely packed
✓ Visual pigment consists of an opsin protein
✓ Internal colour representation of the external world -
spectral sensitivity and spatial distribution of
photoreceptors as well as screening pigments
(Song and Lee, 2018)
T Y P E S
O F
P H O T O R E C E P T O R S
NUMBER OF
PHOTORECEPTOR
INSECT NAME PHOTORECEPTOR REFERENCE
2 Periplaneta americana UV and green Honkanen et
al., 2014
3 Most insects UV, blue, and green Briscoe and
Chittka, 2001
4 Odonata and Lepidoptera Red, UV, green, and blue-
photoreceptors
Peitsch et al.,
1992
5 Butterfly (Papilio xuthus) UV, violet, narrow blue, single
green and Red
Kelber et al.,
2003
Dragonfly (Sympetrum
rubicundulum)
Red housefly (Musca
domestica)
6
Spectral receptor
classes
Bluebottle butterfly
(Graphium sarpedon
nipponum)
UV,violet,blue, blue–green,
green and red
(15 spectral sensitivities)
Chen et al.,
2016
8 Drosophila melanogaster Six outer photoreceptors and
Two inner photoreceptors
Katz and
Minke, 2009
TAPETUM
❖ Reflective layer located behind the retina
in their eyes, which acts like a mirror to
bounce light back through the photoreceptor
cells, enhancing their vision in low-light
conditions
❖ Present in moths and butterflies
❖ Some butterflies may have a less
developed tapetum
❖ They evolved to have less need for such a
reflective layer due to their daytime
activity
(Takemura et al., 2007)
IMAGE FORMATION
✓ The ommatidia are packed side by side – Wide field of view
✓ Ommatidia is oriented in a slightly different direction – Honey
comb
✓ Each ommatidium produces an image of a small part of the object
✓ Small images – Combined in brain - complete image - Mosaic
picture
✓ Image lacks depth - poor in resolving - excellent at detecting
movement
✓ High flicker fusion rate - Action of an object -independent
frames
✓ Picking up motion of the object - Escape predators and catch prey
(Tomlinson, 2012)
T YPES OF
COMPOUND EYES
TYPES OF COMPOUND EYES
❖ Each ommatidium is isolated from its neighbors -
sleeve of light- absorbing screening pigment
❖ It prevent light - photoreceptive rhabdom
❖ Limiting light capture - Lower light sensitivity
❖ Higher image resolution - more detailed image
❖ Diurnal insects like bees, butterflies and dragonflies
(Warrant and Dacke, 2011)
TYPES OF COMPOUND EYES
❖ A large number of corneal facet lenses and
bullet-shaped crystalline cone lenses
❖ They collect and focus light across the clear zone
of the eye - toward single photoreceptor
❖ Several hundred, or even thousands, of facets
service a single photoreceptor
❖ High sensitivity – Low image resolution
❖ Moths , Dung beetle – Nocturnal insects
f - focal length
c - corneal facet
lens
cc - crystalline
cones
p - screening
pigment
rh - rhabdom
cz - clear zone
(Warrant and Dacke, 2011)
COLOUR VISION IN INSECTS
✓ Light spectrum - rich information about the world
✓ Color vision - internal representation of spectral
distribution in the environment
✓ Colour - important role – foraging – habitat - mates
✓ Photoreceptors - give them the capability to see
colours
✓ Pollinating insects - memorize the colours and
patterns
(Song and lee , 2018)
COLOUR VISION IN
INSECTS
INSECT GROUPS RESPONDED WAVELENGTH
Lepidoptera 360 – 560 nm
Lepidoptera larvae 340-540 nm
Hemiptera 440 nm
Coleoptera 395 nm
(Pan et al., 2021)
DIURNAL INSECT
✓ Insect group that are found to be more
actively abundant during the day
✓ Examples: Bees, wasps, ants, and
thrips
NOCTURNAL INSECT
✓ Insect group that are found to be more
actively abundant during the night
✓ Examples: Caddisflies, moths and
earwigs
CIRCADIAN RHYTHM
❖ It is the body's natural 24-hour cycle that regulates sleep, wakefulness,
and other bodily functions - Internal clock
❖ It controls daily rhythmicity in behavior (locomotion, feeding, mating,
and oviposition) physiological functions, and developmental events
such as hatching, pupation, eclosion etc.,
(Numata et al., 2015)
BENEFITS OF BEING NOCTRUNAL INSECTS
❖ Protection from diurnal predators
❖ Avoid competition for food
(Somanathan et al., 2007)
NOCTRUNAL INSECTS
Earwig Caddishfly
Cockroach
Moth
Click beetle
Glow worm
Austrophasmatidae
Mayfly
NOCTRUNAL INSECTS
Cricket Dung beetle Mosquito Firefly
Bed bug Giant water bug
Green lace wing
Dobsonfly
FUNGUS GNAT
Waitomo cave
❖ Thousands of tiny blue lights coating the
rock surfaces - natural planetarium
❖ Arachnocampa luminosa
❖ Produce light by breaking down a light-
emitting protein
❖ It produce constant glow – for bait
(Kennerson, 2016)
FUNGUS GNAT
❖Worms drop threadlike snares, thin
filaments of silk secreted from their mouths
and dotted with balls of mucous
❖Mucous balls - glass beads, magnify the
worms spectral light – chandeliers
❖ Wind can tangle the sticky strings
❖Deeper inside the cave- longer the
threads
(Kennerson, 2016)
N O C T R U N A L I N S E C T A N D T H E I R R O L E I N E C O S Y S T E M
▪ Pollinator
▪ Decomposer
▪ Predator – prey dynamics
NOCTURNAL POLLINATORS
✓
✓
✓
✓
✓
✓
✓
✓
(Macgregor and Scott, 2020)
Raspy circket (Glomeremus sp)
Formicomus braminus
N O C T U R N A L P O L L I N ATO R S
(Xiong et al., 2020)
Rare Chinese flower (Vincetoxicum hainanense )
Pollinated by
(A) Cecidomyiidae sp
(B) Tetramorium sp
(D) Hexacentrus sp. (Orthoptera)
(E) Carabidae sp
(F) Blattella bisignata
(G) Blattella bisignata
(H) Opisthoplatia orientalis
MECHANISMS FOR NIGHT-TIME POLLINATION
INTERACTIONS
(Wang et al., 2014)
1. FLORAL SCENT 2. NIGHT VISION 3. THERMOGENESIS
Sap beetle,Epuraea sp
Magnolia sprengeri
Petunia sp.
VISUAL NAVIGATION IN NOCTURNAL INSECTS
❖Despite their small eyes and tiny brains,
nocturnal insects have acquired a remarkable
aptitude to visually navigate at night
❖Some utilize - celestial compass cues
❖Others - visual landmarks to navigate
❖Impressive abilities - sensitive compound eyes
and specialized visual processing strategies
❖Extremely difficult to distinguish cues at night -
light levels 11 orders of magnitude lower
(Warrant and Dacke, 2016)
NAVIGATION AND ORIENTATION CUES
❖Insects need accurate detection -
return to the safety of a nest after
foraging trip
❖Effectively escape from competitors
and predators
Celestial cues
Terrestrial cues
(Warrant and Dacke, 2016)
VISUAL NAVIGATION CUES
✓ Moon is a far more complex orientation cue than the sun
✓ Many nocturnal insects use its bright disk for orientation
and navigation
✓ Moon - brightest & visible indicator at night
✓ Moonlight moves toward Earth- scattered by the
atmosphere, circular pattern centered on the moon
✓ Moon's polarization pattern, which is a million times dimmer
✓ Scarabaeus zambesianus - nocturnal dung beetle -
straight line
(Freas and Spetch, 2023)
POLARIZATION OF
LIGHT
VISUAL NAVIGATION CUES
✓ Many nocturnal insects - rely on landmarks for
orientation, flight control and to locating a nest entrance
in the dark.
✓ Contrast against the background than their own
luminance
✓ Gypsy moths and mosquitoes use optic flow signals to
control their flight paths in low light
(Freas and Spetch, 2023)
C L A S S E S O F N O C T U R N A L N AV I G AT I O N I N I N S E C T S
❖
❖
❖
(Warrant and Dacke, 2016)
S H O R T D I S TA N C E S T R A I G H T- L I N E O R I E N TAT I O N
➢ Scarabaeus satyrus and S. zambezianus beetles
search for fresh dung after the sun sets – South
Africa
➢ Safest and most effective way to leave this
battlefield - straight as possible
➢ Principal orientation cue- Celestial pattern of
polarized light –moon
➢ Milky Way – when moon is deep below the horizon
(Warrant and Dacke, 2016)
DUNG BEETLE – MILKYWAY ORIENTATION
➢ When the moon is absent from the night sky,
stars remain as celestial visual cues
➢ In a planetarium, the beetles orientate
equally well when rolling under a full starlit
sky as when only the Milky Way is present
➢ First documented use of the Milky Way
for orientation in the animal kingdom
(Dacke et al., 2013)
LONG-DISTANCE HIGH -ALTITUDE MIGRATION
❖ Australian Bogong moth Agrotis infusa, perform spectacular long-distance seasonal migrations.
❖ 650 kilometres in a single night
❖ Radar recording reports – moth activity in Moon less nights – Milky way as a cue
❖ Nocturnal moths might have a magnetic compass sense to help them maintain their migratory
course
(Warrant and Dacke, 2016)
HOMING
❖ In order to get home, regardless of the time of day, an
animal needs to know where it is in relation to a goal
❖ Many bits of information are usually used to calculate
this present position
❖ Path integration - length and direction of its “home
vector”
❖ Using visual landmarks and integrating paths are
crucial tactics
(Warrant and Dacke, 2016)
NIGHT VISION AND INSECT PHOTORECEPTORS
➢ Photons – Short supply - difficult to visually guide behavior
➢ Photoreceptors that effectively transform photon neural
signal
➢ Photon absorption rate - Periplaneta photoreceptors is 0.1
photons per second- 17 times higher in flies, 50 times
higher in nocturnal bees like Megalopta genalis, and
4400 times higher in hawkmoths like Deilephila
➢ Not imply that these insects require more ambient light to
see
➢ Megalopta and Periplaneta have apposition eyes -
nocturnal - larger lens and rhabdom diameter – Anatomical
changes
(Honkanen et al., 2017)
➢ Insect photoreceptors/microvillar - Thousands of bristle-like microvilli
➢ Optical pigment rhodopsin incorporated in the microvillar membrane -
photons that enter rhabdom
➢ Phototransduction cascade - Quantum bump
➢ Nocturnal insect photoreceptors – large quantum bumps
➢ Example - Tropical sweat bees Lasioglossum and Megalopta
➢ Cockroaches (Heimonen et al., 2012), locusts (Lillywhite, 1977),
carpenter ants (Souza et al., 1989), crane flies (Laughlin and
Weckstrom, 1993), and stick insects (Frolov et al., 2012)
NIGHT VISION AND INSECT PHOTORECEPTORS
(Honkanen et al., 2017)
COLOUR VISION IN NOCTURNAL INSECTS
➢ Human observer nocturnal world - Black and white
➢ Day to night - three spectral classes photoreceptors—world
with colour—begin to fail
(Kelber and Lind, 2010)
➢ The physical colour of an object
Irradiance spectrum of natural daylight
Spectral reflectance properties of the object’s surface
(Warrant and Somanathan , 2022)
COLOUR VISION IN NOCTURNAL INSECTS
Advantages of colour vision
➢ Recognizing food
➢ Mates
➢ Habitats
(Somanathan et al., 2008; Kelber et al., 2003)
PROBLEMS IN COLOUR VISION AT DIM LIGHT
PROBLEM 1
❑ Colours and contrasts of the visual world are same at night
❑ 100 billion times dimmer than sunshine - discrimination
unreliable
PROBLEM 2
❑ Visual noise – Photoreceptor uncertainty of photon
arrival
❑ Photon shot noise - reduce reliability of intensity
discrimination -contrast details
(Warrant, 2017)
SIGNAL TO NOISE RATIO
❑ SNR - Improves with increasing photon catch - worse
at lower light levels
❑ ‘de Vries-Rose’ or ‘square root law’ of visual detection at
low light levels indicates that the visual SNR, and thus
contrast discrimination, improves as the square-root of
photon catch
𝑁
√𝑁
(Warrant and Somanathan, 2022)
TRANSDUCER NOISE
➢
➢
DARK NOISE
➢ Biochemical pathways - transduction are
occasionally activated - even in perfect
darkness - thermal activations of
rhodopsin molecules
➢ Continuous low amplitude fluctuation
➢ Electrical responses that are
indistinguishable from those produced by real
photons
➢
TYPES OF NOISE
( )
COLOUR VISION
VS
NOISE
❖ Dimmer light levels and higher noise levels,
fewer colours can be seen
❖ Nocturnal colour vision is rare in vertebrates
❖ Photoreceptors of vertebrates are generally
much noisier than those of insects
❖ Monochromatic vision default at night
(Warrant and Somanathan, 2022)
HOW INSECT FIND SOLUTION TO
COLOUR VISION
➢ Discriminating colour in dim light - low visual SNR
➢ Solution lies in optical and neural strategies
➢ Increasing the photon catch or Reducing the noise
➢ Insects have evolved many such strategies
o Optically - signal amplitude
can be improved by having
an eye design that captures
more light
o Peripheral neural
mechanisms that sum
photons of light in time and
space
(Warrant and Somanathan, 2022)
SUMMATION
SPATIAL SUMMATION
❑ Instead of each channel collecting
photons
❑ Activation of specialized laterally
spreading neuron - couple visual
channels into groups
TEMPORAL SUMMATION
❑ Increasing longer exposure time
❑ Responding more slowly and
building up a brighter image
❑ Fast-moving object, can be
significantly degraded
➢ Summation improve the visual SNR in dim light
➢ Coarser and slower features of the world at the expense of the finer and faster
features.
➢ Absence of summation nothing seen
(Laughlin and Weckstrom, 1993)
A L G O R I T H M I N S P I R E D B Y N O C T U R N A L V I S U A L
P R O C E S S I N G
❑ Amplification of primary image
signals to optimized spatio–
temporal summation to reduce
noise
❑ Increases the reliability of video
collected in dim light, including
the preservation of color
(Warrant et al., 2014)
HAWK MOTH NOCTURNAL COLOUR VISION
❖ Most hawkmoth are active at dawn and dusk
❖ To find and recognize rewarding flowers independent of
the colour of the illuminating light – colour constant
visual system
❖ Behavioural experiments shows that the nocturnal
hawkmoth Deilephila elpenor uses colour vision to
discriminate coloured stimuli at intensities
corresponding to dim starlight (0.0001 cd m-2)
❖ First report for nocturnal colour vision in animals
(Kelber et al., 2002)
NOCTURNAL COLOUR VISION
❖ Generally assumed nocturnal flowers attract insects by
their high intensity contrast to the back ground of green
leaves
❖ White nocturnal flowers absorb ultraviolet light
❖ Rhodopsins have some sensitivity in the ultraviolet range
❖ Flowers are therefore not maximally bright to an insect
eye
(Warrant and Somanathan, 2022)
E X P E R I M E N T O N
CO LO U R V I S I O N
❖ Trained D. elpenor to associate a
reward of sugar solution with a
colour, either blue or yellow to a
human observer
❖ Five different light intensities ranging
from mid-dusk (1cdm-2) to dim
starlight (0.0001cdm-2)
(Kelber et al., 2002)
E X P E R I M E N T O N CO LO U R V I S I O N
Choice frequencies in tests after
training to blue between
humans and D. elpenor
Choice frequencies in tests after
training to yellow between
humans and D. elpenor
(Kelber et al., 2002)
ILLUMINATION AND COLOUR
CONSTANCY
❑ Discrimination was tested in two
different illumination colours, white
and broad-spectrum yellow
❑ Without colour constancy moths
would confuse these two stimuli
Moths trained to green stimuli (b) and turquoise stimuli
(c) chose correctly in tests under two different
illuminations, `white' (left) and `yellow' (right)
(Kelber et al., 2002)
TWO TYPES OF OMMATIDIA
✓ D. elpenor has two different types of
ommatidium in the eye: Both types have seven
proximal green receptors - one has two distal
blue receptors, and other has two distal
ultraviolet receptors.
✓ Quantum catches for the ultraviolet
photoreceptor were especially low.
✓ 0.0001 cd/m2
✓ Photon shot noise more
✓ Reliable discrimination of blue from grey is
impossible under these conditions - spatial and
temporal summation
Quantum catches - D. elpenor photoreceptors
(Kelber et al., 2002)
COLOUR
VISION IN BEES
❖ Colour vision in the Apis mellifera -
model
❖ Three spectral classes of
photoreceptors - ultraviolet (344 nm),
blue (436 nm), and green (556 nm)
portions of the spectrum enable colour
vision
❖ Colour perception reduced in light
levels
(Vijayan et al., 2023)
COLOUR VISION IN ROCK BEE
Nocturnal foraging in A. dorsata - lunar phase - third quarter maximum activity
❖ Colour vision thresholds of
nocturnally active insects
❖ The violet, blue and green bands
represent trichromacy in these
species
(Vijayan et al., 2023)
EXPERIMENT ON
COLOUR VISION
IN ROCK BEE
(Vijayan et al., 2023)
MEGALOPTA
GENALIS
✓ Sweat bee Megalopta (Hymenoptera:
Halictidae)
✓ Nocturnal behaviour is a successful strategy
✓ Megalopta performs visually guided flight with
apposition eyes
✓ Forage only when the sun is down: shortly
before sunrise, and shortly after sunset
(Warrant et al., 2004)
✓ Megalopta nests are often found under thick
canopy in the understory dim in bright daylight.
❖ Female Megalopta genalis have relatively
large eyes
❖ Corneal facet diameters are 1.8 times
larger, and rhabdom diameters 4–5 times
larger, than those of diurnal halictids
(Greiner et al., 2004)
❖ These optical adaptations together make
the eyes of M. genalis 27 times more
sensitive than those of their diurnal
counterparts
MEGALOPTA GENALIS
MEGALOPTA VS LASIOGLOSSUM
❖ (a, b) Responses to single photons
❖ Bump amplitude is larger, and the bump time
course much slower, in M. genalis than in L.
leucozonium
❖ In light-adapted conditions (c, d), both species reach
the same maximum contrast gain per unit bandwidth,
although L. leucozonium - broader bandwidth
❖ In dark-adapted conditions (e, f), M. genalis has a
much higher contrast gain per unit bandwidth
photoreceptor responses
❖ Larger bumps of M. genalis - Photoreceptor’s gain of
transduction is greater compared with diurnal species
Large bumps in nocturnal crane flies (Stavenga et al.,
1993), cockroaches (Greiner et al., 2004) ,bees (Matsuura,
1999)
NEURAL STRATERGY
❑ Spatial summation, which improves visual
reliability by grouping signals from
neighbouring photoreceptors
❑ Temporal summation, which does so by
increasing integration times
❑ Otherwise plagued by photon noise
❑ Trade resolution for sensitivity is if it improves
visual performance
(Snyder et al., 1977)
LOW LIGHT LEVELS AND MEGALOPTA
FLIGHT PERFORMANCE
✓ Bright light landings were always quick
✓ Low light landings included both quick
and slow samples
✓ Nocturnal hovering insect's hawkmoths,
lose acuity to spatial pooling
✓ Even as light levels and acuity drop,
bees cannot slow their approaches
✓ Bees learn the speed at which landmark
images move across the retina - goal
(Cartwright and Collett, 1979)
E X P E R I M E N T O N M E G A L O P TA –
L A N D M A R K O R I E N TAT I O N
✓ Orientation flight - Spatial
arrangement of landmarks around the
nest entrance
✓ Five nests in a row, about 1 m above
the forest floor
✓ Nest Leaving light intensity – 0.01
cd/m2
✓ Returning intensity - 0.0001 cd/m2
(Zeil et al., 1996)
Two landmark-manipulation experiment
OPTICAL PARAMETER IN MEGALOPTA
N=1.13(
𝝅
𝟒
)∆p2 D2 k𝝉
∆
t‫׬‬ 𝟏 − 𝒆−𝑲𝑹 𝝀 𝒍
/( 𝝀)𝒅𝝀
(Souza et al.,1989)
✓ Megalopta eyes have morphological, optical, and electrophysiological
characteristics - suit to nocturnal life
✓ In Megalopta 0.15 photons are absorbed by a single green receptor -
one integration time
✓ In Apis at the same intensity, N = 0.0053 photons
Acceptance angle -
rhabdomere - range of
angles from which it
can effectively absorb
light
N O C T R U N A L C A R P E N T E R
B E E
✓ Xylocopa tranquebarica nocturnal activity
on moonlit nights
(Burgett and Sukumalanand, 2000)
✓ Xylocopa tenuiscapa and Xylocopa
leucothorax are potential competitors
✓ Exploit that flower for pollen and nectar
rewards at night
(Hopkins et al., 2000)
F L I G H T A C T I V I T Y O F T H R E E S P E C I E S O F C A R P E N T E R B E E S
(Somanathan et al., 2007)
L A N D M A R K O R I E N TAT I O N E X P E R I M E N T I N X Y L O C O PA
(Somanathan et al., 2007)
(Somanathan et al., 2007)
ANATOMY OF NOCTRUNAL CARPENTER BEE EYES
Optical sensitivities - X. tranquebarica has eyes that are 27 times more sensitive
than X. leucothorax and 9 times more sensitive than those of X. tenuiscapa
OPTICAL
SENSITIVITY
OCELLI OF THREE CARPENTER BEE
X. leucothorax (a, d)
X. tenuiscapa (b, e)
X. tranquebarica (c, f)
Tracheal tapetum in the ocelli of X. tranquebarica
others instead have dark pigmentation below the ocellar retina
(Somanathan et al., 2009)
PATH INTEGRATION
✓ Path integration - Cataglyphis ants
✓ Path integration - leaving a starting point, such as a nest, an animal
updates an accumulator that keeps a running tally of its current direction
and distance - path back to starting point
(Collett and Collett, 2000)
N AV I G AT I O N U S I N G C A N O P Y PAT T E R N
❖ Parastrachia japonensis (Heteroptera: Parastrachiidae)
❖ Female leaves her burrow to find drupes - Schoepfia jasminodora (Olacaceae:
Rosidae: Santales)
❖ It inserts the proboscis into the drupe and drags - burrow
❖ Direct celestial cues are likely to be unreliable – canopy pattern used
Sudden 180◦ rotation of an artificial
canopy gap in the roof of its box - 12 bugs
(Hironaka et al., 2008)
NOCTURNAL BULL ANTS AND ITS NAVIGATION
Study revealed that nocturnal bull ants
(Myrmecia midas) can navigate using
the very faint polarization pattern
produced by moonlight
(Freas et al., 2024)
NOCTURNAL INSECT MIGRATION
➢ Europe’s death’s-head hawkmoths
(Acherontia Atropos)
➢ 2,000 miles from Europe to Africa
➢ 14 hawkmoths and recorded their precise
GPS locations from a light aircraft -
33.8km/hour
➢ Combination of visual landmarks and
Earth’s magnetic field + Favorable tail
winds
(Menz et al., 2022)
NOCTURNAL PEST
➢ Many insects are nocturnal pest, including bed
bugs, earwigs, moths, cockroaches etc.,
➢ According to research, Fall Armyworm outbreak
could potentially lead to maize production losses
ranging from 4.1 to 17.7 mil.tons/year 12 maize-
producing countries in Sub-Saharan Africa
(Rwomushana et al., 2018)
➢ Mosquito-borne diseases cause significant
economic damage globally, costing billions of
dollars each year
LIGHT TRAP
➢ Insects see ultraviolet (UV) radiation
➢ Nocturnal insects are often attracted to light
sources that emit large amounts of UV radiation
➢ Devices that exploit this behavior, such as light
traps for forecasting pest outbreaks, and electric
insect killers – developed
➢ Lamps - yellow illumination effectively to control
the activity of nocturnal moths
(Shimoda and Honda , 2013)
YELLOW FLUORESCENT LAMPS
(Shimoda and Honda , 2013)
Flower cultivation facilities
with yellow fluorescent
lamps
A: Carnation
B: Chrysanthemum
LEDS IN PEST CONTROL
❖ Electrical current passes through a microchip - emit
light
❖ Different colors - semiconductor materials that emit
photons at different wavelengths
❖ Wide spectrum of colours - red, green, and blue
(RGB) light
ADVANTAGES
✓ Species-specific response
✓ Precise wavelength control
✓ Improved attraction rates
✓ Reduced energy
consumption
✓ Heat reduction
(Wakefield et al., 2016)
NOCTURN AL INSECT MONITORING METHODS
(Roy et al., 2024)
❑ Search Light trap
❑ Vertical-looking entomological radar
❑ Camera trap
The IMR (Insect Monitoring Radar) Tower,
designed by Dr. Christopher M. Kaltenbach,
is inspired by the UAE’s historic
watchtowers
NOCTURN AL INSECT MONITORING METHODS
(Roy et al., 2024)
AUTOMATED SENSOR
❑ Insects - high diversity - 80%
❑ Sensors comprise a light to attract
insects, a camera for collecting images
and a computer for scheduling, data
storage and processing
❑ Metadata is important - balance the
capture - power and data storage
limitations
NOCTURN AL INSECT MONITORING METHODS
(Sun et al., 2022)
SEARCH LIGHT TRAP
❑ Searchlight trapping is a method that
uses light to attract and trap migratory
insects
❑ Three species of rice migratory pests
(Cnaphalocrocis medinalis, Sogatella
furcifera, and Nilaparvata lugens)
cause severe yield and economic losses
to rice food every year in china
NOCTURN AL INSECT MONITORING METHODS
(Roy et al., 2024)
Machine learning workflow to analyse moth camera trap data
NOCTURN AL INSECT MONITORING METHODS
(Cheng et al., 2018; Gao et al., 2024)
RADAR
❑ Migration is a key process in the numerous
insect species – Pests
❑ Identification of insect migrants is important
❑ Vertical-looking radars (VLRs) - monitor high-
altitude insect migration
❑ The VLRs emit a vertically pointing, linear-
polarized, narrow-angle conical scan
❑ Insect migrate in tightly clustered – bright moon
nights – clouds were sparse
IMPACT OF ARTIFICIAL
LIGHT AT NIGHT ❖ Localized illumination of nocturnal
landscapes by anthropogenic sources
of light
❖ Global surveys - 8% above the natural
level
(Falchi et al., 2016)
❖ Insect diversity and abundance - rapid
decline
(Hallmann et al., 2017)
❖ In Nocturnal insects - spatial
disorientation and attraction
❖ Fireflies ,Click beetles and Glowworms -
vulnerable to artificial illumination
TEMPORAL DISORIENTATION
❖ ALAN - temporal disorientation, desynchronization of
organisms from their biorhythms
(Saunders, 2012)
❖ Insect - circadian patterns of activity (foraging,
reproduction, migration, etc.) - light cycle
❖ Nocturnal insects - emergence time , feeding and
courtship - dictated by internal clocks - ambient light
(Saunders, 2009)
DESENSITISATION
❖ Nocturnal insects - highly sensitive visual systems
– not function well in illuminated environment
❖ Some insects like Myrmecia ants are capable of
flexible, rapid light adaptation
(Narendra et al., 2013)
❖ Many photons at once, some insects - temporarily
dazzled or permanently blinded
❖ Gryllus bimaculatus - exposure to bright UV light
structural degeneration of photoreceptors
ALAN ON
BIOLUMINESCENT
INSECTS
✓ 2000 lampyrid species
✓ Adult fireflies - employ bioluminescence -
courtship signal
✓ Click beetles (Coleoptera: Elateridae),
Railroad worm (Coleoptera: Phengodidae),
and Fungus gnat (Diptera: Keroplatidae),
employ bioluminescence as an aposematic
signal or predatory lure
(Rochow, 2007)
DORSAL LIGHT RESPONSE
 Interaction between flying insects
and artificial light – “Drawn like a
moth to a flame”
 The dorsal-light-response is a
behavior where flying insects keep
their top side the dorsal side
pointed at the brightest area in
their vision
 Artificial light is an ancient method -
Roman Empire - 1 AD
Three flight patterns
1.Orbiting: a circular flight around the bulb - body
slightly tilted to the light.
2. Stalling: a sharp climb away from the ground as
the insect faces away from the light source
3.Inversion: a dive to the ground
(Owens and Lewis, 2018)
WHY INSECTS DRAWN TO LIGHT ?
Insects use the moon as a celestial compass cue to navigate - artificial light sources
Thermal radiation from light sources is attractive to flying insects
Insects are drawn to light through an escape mechanism -gap in the foliage
(Owens and Lewis, 2018)
SOLUTIONS TO REDUCE LIGHT POLLUTION
❖ Avoid blue light Insects are particularly attracted to
blue wavelengths
❖ Red light It has minimal impact on insects
❖ Shielding: Install fixtures with hoods or baffles to
direct light downwards
(Schroer et al., 2021)
SOLUTIONS TO REDUCE LIGHT POLLUTION
❖ Only light when needed: Turn off lights not in use
❖ Motion sensors: Illuminate when movement
detected
❖ Dimmers: Reduce light intensity when necessary
❖ Awareness campaigns: Educate your community
about the impacts of light pollution on insects
(Schroer et al., 2021)
BIOMIMICRY INSECT COMPOUND EYE
BIOMIMICRY INSECT COMPOUND EYE
CONCLUSION
❑ Nocturnal insects play the same
important role as diurnal insects
❑ They have successfully adapted to the
darkest environments
❑ Vision plays a major role
❑ “By reducing light pollution we can
save the unseen beauty of night world”
THANK YOU

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Vision and visual navigation in nocturnal insects

  • 1. VISION AND VISUAL NAVIGATION IN NOCTURNAL INSECTS PRESENTED BY T.YUVASRI 23PGA205 CHAIRMAN : Dr. M. KANDIBANE PANDIT JAWAHARLAL NEHRU COLLEGE OF AGRICULTURE AND RESEARCH INSTITUTE, KARAIKAL- 609603
  • 2. INTRODUCTION ❖ Insects play tremendous - ecological and economic role globally ❖ Insect activity is on average 31.4% higher at night than in the day (Wong and Didham, 2024) ❖ Insects have extremely complex vision systems - small eyes and relatively tiny brains (Warrant, 2017)
  • 3. INSECT EYES ❖ Insect eyes are a marvel of evolutionary adaptation ❖ Unlike the simple, single lens eyes - vertebrates, Insects possess compound eyes ❖ Insect eyes are adapted to various ecological niches-specialized visual systems to suit their specific needs (Buschbeck and Friedrich, 2008)
  • 4. INSECT EYES EVOLUTION ❖The origin of compound eyes can be traced back to the Cambrian period - 500 million years ago ❖Modern insect ancestor eyes only detect light and dark ❖Simple eyespots to compound eyes involved incremental changes that provided - Better predator detection and improved navigation (Warrant and Nilsson, 2006) DORSAL OCELLI
  • 6. STRUCTURE OF INSECT EYES LATERAL OCELLI DORSAL OCELLI COMPOUND EYE SIMPLE EYE TYPES
  • 7. SIMPLE EYES ❖ It comprise of single lens for collecting and focusing light SIMPLE EYES LATERAL OCELLI DORSAL OCELLI
  • 8. LATERAL OCELLI ❖ Larval vision relies on single-lensed eyes called stemmata ❖ Larval hymenopterans (excluding symphytans) and fleas do not possess stemmata (Simpson and Douglas, 2013) ❖ Sawflies and Tiger beetles have more advanced stemmata with improved visual abilities (Paulus, 1986)
  • 9. STEMMATA OF FIREFLY ❖ Photuris larvae - single stemma with each eye containing 88 photoreceptors ❖ Fusion stemmata - Two clusters of cells fuse to form the dorsal stemma, and three ventral clusters merge to form the ventral stemma ❖ Wide, net-like rhabdomere - vertical columns shared central region - each rhabdom has a ‘fused’ organization - consistent with rhabdoms -maximal photon capture (Murphy and Moiseff, 2019)
  • 10. DORSAL OCELLI ✓ Dorsal ocelli - front surface of the head of nymphs and adults of several insect ✓ Present in Bees, locusts and dragonflies ✓ It cannot form an image and is unable to recognize the object ✓ Sensors of ambient light intensity e.g., for the control of circadian rhythms (Honkanen et al., 2018)
  • 11. COMPOUND EYE ❖ Compound eyes are the primary visual receptors of adult insects and larval hemimetabola ❖ They are constructed from many similar units called ommatidia Lepismatidae - 12 ommatidia on each side Collembola - 8 widely spaced ommatidia Dragonfly - 30,000 Drone bees - 10,000 Ponera punctatissima - 1 (Simpson and Douglas, 2013)
  • 12. STRUCTURE OF COMPOUND EYE OMMATIDIA CORNEAL LENS CRYSTALLINE CONE PRIMARY PIGMENT CELLS SECONDARY PIGMENT CELLS RHABDOM PHOTORECEPTOR BASEMENT MEMBRANE
  • 13. PARTS OF COMPOUND EYE OMMATIDIA Light-gathering part and a sensory part, perceiving the radiation and transforming it into electrical energy CORNEAL LENS ❖ The cuticle covering the eye is transparent and colorless - covers outer ommatidium ❖ It is produced by corneagen cells ❖ Later withdraw to the sides primary pigment cells (Zhao et al., 2014) CRYSTALINE CONE ❖ Crystalline cone and the lens work together to refract light onto the rhabdom (Tomlinson, 2012)
  • 14. PARTS OF COMPOUND EYE PIGMENT CELLS ✓ Diurnal insects - pigment cells surrounding each ommatidium – Mosaic image ✓ Nocturnal and crepuscular insects have pigment cells that do not completely isolate each facet less distinct mosaic PRIMARY PIGMENT CELLS ❖ Primary pigment cells in insect eyes create an insulating cylinder around the lens unit, which helps improve visual acuity ❖ Crescent-shaped SECONDARY PIGMENT CELLS ❖ Found in the retina of insect compound eyes - between two adjacent ommatidia ❖ Regulate the amount of light that reaches the photoreceptors (Tomlinson, 2012)
  • 15. PARTS OF COMPOUND EYE RHABDOM ✓ Light−sensitive part ✓ Rod-like structure, secreted by retinula cells and centered on the optical axis ✓ It contains light-sensitive pigments – rhodopsin ✓ Pigments absorbs wavelength of incident light and generate nerve impulses - photochemical process ✓ Resolve wavelength and plane of polarization Fused rhabdom – Bees Open rhabdom - Fruitflies (Zelhof et al., 2006)
  • 16. PARTS OF COMPOUND EYE PHOTORECEPTOR ✓ Ommatidium - eight to nine photoreceptors ✓ Photoreceptors have rhabdomeres, microvillar structures where visual pigments are densely packed ✓ Visual pigment consists of an opsin protein ✓ Internal colour representation of the external world - spectral sensitivity and spatial distribution of photoreceptors as well as screening pigments (Song and Lee, 2018)
  • 17. T Y P E S O F P H O T O R E C E P T O R S NUMBER OF PHOTORECEPTOR INSECT NAME PHOTORECEPTOR REFERENCE 2 Periplaneta americana UV and green Honkanen et al., 2014 3 Most insects UV, blue, and green Briscoe and Chittka, 2001 4 Odonata and Lepidoptera Red, UV, green, and blue- photoreceptors Peitsch et al., 1992 5 Butterfly (Papilio xuthus) UV, violet, narrow blue, single green and Red Kelber et al., 2003 Dragonfly (Sympetrum rubicundulum) Red housefly (Musca domestica) 6 Spectral receptor classes Bluebottle butterfly (Graphium sarpedon nipponum) UV,violet,blue, blue–green, green and red (15 spectral sensitivities) Chen et al., 2016 8 Drosophila melanogaster Six outer photoreceptors and Two inner photoreceptors Katz and Minke, 2009
  • 18. TAPETUM ❖ Reflective layer located behind the retina in their eyes, which acts like a mirror to bounce light back through the photoreceptor cells, enhancing their vision in low-light conditions ❖ Present in moths and butterflies ❖ Some butterflies may have a less developed tapetum ❖ They evolved to have less need for such a reflective layer due to their daytime activity (Takemura et al., 2007)
  • 19. IMAGE FORMATION ✓ The ommatidia are packed side by side – Wide field of view ✓ Ommatidia is oriented in a slightly different direction – Honey comb ✓ Each ommatidium produces an image of a small part of the object ✓ Small images – Combined in brain - complete image - Mosaic picture ✓ Image lacks depth - poor in resolving - excellent at detecting movement ✓ High flicker fusion rate - Action of an object -independent frames ✓ Picking up motion of the object - Escape predators and catch prey (Tomlinson, 2012)
  • 21. TYPES OF COMPOUND EYES ❖ Each ommatidium is isolated from its neighbors - sleeve of light- absorbing screening pigment ❖ It prevent light - photoreceptive rhabdom ❖ Limiting light capture - Lower light sensitivity ❖ Higher image resolution - more detailed image ❖ Diurnal insects like bees, butterflies and dragonflies (Warrant and Dacke, 2011)
  • 22. TYPES OF COMPOUND EYES ❖ A large number of corneal facet lenses and bullet-shaped crystalline cone lenses ❖ They collect and focus light across the clear zone of the eye - toward single photoreceptor ❖ Several hundred, or even thousands, of facets service a single photoreceptor ❖ High sensitivity – Low image resolution ❖ Moths , Dung beetle – Nocturnal insects f - focal length c - corneal facet lens cc - crystalline cones p - screening pigment rh - rhabdom cz - clear zone (Warrant and Dacke, 2011)
  • 23. COLOUR VISION IN INSECTS ✓ Light spectrum - rich information about the world ✓ Color vision - internal representation of spectral distribution in the environment ✓ Colour - important role – foraging – habitat - mates ✓ Photoreceptors - give them the capability to see colours ✓ Pollinating insects - memorize the colours and patterns (Song and lee , 2018)
  • 24. COLOUR VISION IN INSECTS INSECT GROUPS RESPONDED WAVELENGTH Lepidoptera 360 – 560 nm Lepidoptera larvae 340-540 nm Hemiptera 440 nm Coleoptera 395 nm (Pan et al., 2021)
  • 25. DIURNAL INSECT ✓ Insect group that are found to be more actively abundant during the day ✓ Examples: Bees, wasps, ants, and thrips NOCTURNAL INSECT ✓ Insect group that are found to be more actively abundant during the night ✓ Examples: Caddisflies, moths and earwigs
  • 26. CIRCADIAN RHYTHM ❖ It is the body's natural 24-hour cycle that regulates sleep, wakefulness, and other bodily functions - Internal clock ❖ It controls daily rhythmicity in behavior (locomotion, feeding, mating, and oviposition) physiological functions, and developmental events such as hatching, pupation, eclosion etc., (Numata et al., 2015)
  • 27. BENEFITS OF BEING NOCTRUNAL INSECTS ❖ Protection from diurnal predators ❖ Avoid competition for food (Somanathan et al., 2007)
  • 28. NOCTRUNAL INSECTS Earwig Caddishfly Cockroach Moth Click beetle Glow worm Austrophasmatidae Mayfly
  • 29. NOCTRUNAL INSECTS Cricket Dung beetle Mosquito Firefly Bed bug Giant water bug Green lace wing Dobsonfly
  • 30. FUNGUS GNAT Waitomo cave ❖ Thousands of tiny blue lights coating the rock surfaces - natural planetarium ❖ Arachnocampa luminosa ❖ Produce light by breaking down a light- emitting protein ❖ It produce constant glow – for bait (Kennerson, 2016)
  • 31. FUNGUS GNAT ❖Worms drop threadlike snares, thin filaments of silk secreted from their mouths and dotted with balls of mucous ❖Mucous balls - glass beads, magnify the worms spectral light – chandeliers ❖ Wind can tangle the sticky strings ❖Deeper inside the cave- longer the threads (Kennerson, 2016)
  • 32. N O C T R U N A L I N S E C T A N D T H E I R R O L E I N E C O S Y S T E M ▪ Pollinator ▪ Decomposer ▪ Predator – prey dynamics
  • 33. NOCTURNAL POLLINATORS ✓ ✓ ✓ ✓ ✓ ✓ ✓ ✓ (Macgregor and Scott, 2020) Raspy circket (Glomeremus sp) Formicomus braminus
  • 34. N O C T U R N A L P O L L I N ATO R S (Xiong et al., 2020) Rare Chinese flower (Vincetoxicum hainanense ) Pollinated by (A) Cecidomyiidae sp (B) Tetramorium sp (D) Hexacentrus sp. (Orthoptera) (E) Carabidae sp (F) Blattella bisignata (G) Blattella bisignata (H) Opisthoplatia orientalis
  • 35. MECHANISMS FOR NIGHT-TIME POLLINATION INTERACTIONS (Wang et al., 2014) 1. FLORAL SCENT 2. NIGHT VISION 3. THERMOGENESIS Sap beetle,Epuraea sp Magnolia sprengeri Petunia sp.
  • 36. VISUAL NAVIGATION IN NOCTURNAL INSECTS ❖Despite their small eyes and tiny brains, nocturnal insects have acquired a remarkable aptitude to visually navigate at night ❖Some utilize - celestial compass cues ❖Others - visual landmarks to navigate ❖Impressive abilities - sensitive compound eyes and specialized visual processing strategies ❖Extremely difficult to distinguish cues at night - light levels 11 orders of magnitude lower (Warrant and Dacke, 2016)
  • 37. NAVIGATION AND ORIENTATION CUES ❖Insects need accurate detection - return to the safety of a nest after foraging trip ❖Effectively escape from competitors and predators Celestial cues Terrestrial cues (Warrant and Dacke, 2016)
  • 38. VISUAL NAVIGATION CUES ✓ Moon is a far more complex orientation cue than the sun ✓ Many nocturnal insects use its bright disk for orientation and navigation ✓ Moon - brightest & visible indicator at night ✓ Moonlight moves toward Earth- scattered by the atmosphere, circular pattern centered on the moon ✓ Moon's polarization pattern, which is a million times dimmer ✓ Scarabaeus zambesianus - nocturnal dung beetle - straight line (Freas and Spetch, 2023)
  • 40. VISUAL NAVIGATION CUES ✓ Many nocturnal insects - rely on landmarks for orientation, flight control and to locating a nest entrance in the dark. ✓ Contrast against the background than their own luminance ✓ Gypsy moths and mosquitoes use optic flow signals to control their flight paths in low light (Freas and Spetch, 2023)
  • 41. C L A S S E S O F N O C T U R N A L N AV I G AT I O N I N I N S E C T S ❖ ❖ ❖ (Warrant and Dacke, 2016)
  • 42. S H O R T D I S TA N C E S T R A I G H T- L I N E O R I E N TAT I O N ➢ Scarabaeus satyrus and S. zambezianus beetles search for fresh dung after the sun sets – South Africa ➢ Safest and most effective way to leave this battlefield - straight as possible ➢ Principal orientation cue- Celestial pattern of polarized light –moon ➢ Milky Way – when moon is deep below the horizon (Warrant and Dacke, 2016)
  • 43. DUNG BEETLE – MILKYWAY ORIENTATION ➢ When the moon is absent from the night sky, stars remain as celestial visual cues ➢ In a planetarium, the beetles orientate equally well when rolling under a full starlit sky as when only the Milky Way is present ➢ First documented use of the Milky Way for orientation in the animal kingdom (Dacke et al., 2013)
  • 44. LONG-DISTANCE HIGH -ALTITUDE MIGRATION ❖ Australian Bogong moth Agrotis infusa, perform spectacular long-distance seasonal migrations. ❖ 650 kilometres in a single night ❖ Radar recording reports – moth activity in Moon less nights – Milky way as a cue ❖ Nocturnal moths might have a magnetic compass sense to help them maintain their migratory course (Warrant and Dacke, 2016)
  • 45. HOMING ❖ In order to get home, regardless of the time of day, an animal needs to know where it is in relation to a goal ❖ Many bits of information are usually used to calculate this present position ❖ Path integration - length and direction of its “home vector” ❖ Using visual landmarks and integrating paths are crucial tactics (Warrant and Dacke, 2016)
  • 46. NIGHT VISION AND INSECT PHOTORECEPTORS ➢ Photons – Short supply - difficult to visually guide behavior ➢ Photoreceptors that effectively transform photon neural signal ➢ Photon absorption rate - Periplaneta photoreceptors is 0.1 photons per second- 17 times higher in flies, 50 times higher in nocturnal bees like Megalopta genalis, and 4400 times higher in hawkmoths like Deilephila ➢ Not imply that these insects require more ambient light to see ➢ Megalopta and Periplaneta have apposition eyes - nocturnal - larger lens and rhabdom diameter – Anatomical changes (Honkanen et al., 2017)
  • 47. ➢ Insect photoreceptors/microvillar - Thousands of bristle-like microvilli ➢ Optical pigment rhodopsin incorporated in the microvillar membrane - photons that enter rhabdom ➢ Phototransduction cascade - Quantum bump ➢ Nocturnal insect photoreceptors – large quantum bumps ➢ Example - Tropical sweat bees Lasioglossum and Megalopta ➢ Cockroaches (Heimonen et al., 2012), locusts (Lillywhite, 1977), carpenter ants (Souza et al., 1989), crane flies (Laughlin and Weckstrom, 1993), and stick insects (Frolov et al., 2012) NIGHT VISION AND INSECT PHOTORECEPTORS (Honkanen et al., 2017)
  • 48. COLOUR VISION IN NOCTURNAL INSECTS ➢ Human observer nocturnal world - Black and white ➢ Day to night - three spectral classes photoreceptors—world with colour—begin to fail (Kelber and Lind, 2010) ➢ The physical colour of an object Irradiance spectrum of natural daylight Spectral reflectance properties of the object’s surface (Warrant and Somanathan , 2022)
  • 49. COLOUR VISION IN NOCTURNAL INSECTS Advantages of colour vision ➢ Recognizing food ➢ Mates ➢ Habitats (Somanathan et al., 2008; Kelber et al., 2003)
  • 50. PROBLEMS IN COLOUR VISION AT DIM LIGHT PROBLEM 1 ❑ Colours and contrasts of the visual world are same at night ❑ 100 billion times dimmer than sunshine - discrimination unreliable PROBLEM 2 ❑ Visual noise – Photoreceptor uncertainty of photon arrival ❑ Photon shot noise - reduce reliability of intensity discrimination -contrast details (Warrant, 2017)
  • 51. SIGNAL TO NOISE RATIO ❑ SNR - Improves with increasing photon catch - worse at lower light levels ❑ ‘de Vries-Rose’ or ‘square root law’ of visual detection at low light levels indicates that the visual SNR, and thus contrast discrimination, improves as the square-root of photon catch 𝑁 √𝑁 (Warrant and Somanathan, 2022)
  • 52. TRANSDUCER NOISE ➢ ➢ DARK NOISE ➢ Biochemical pathways - transduction are occasionally activated - even in perfect darkness - thermal activations of rhodopsin molecules ➢ Continuous low amplitude fluctuation ➢ Electrical responses that are indistinguishable from those produced by real photons ➢ TYPES OF NOISE ( )
  • 53. COLOUR VISION VS NOISE ❖ Dimmer light levels and higher noise levels, fewer colours can be seen ❖ Nocturnal colour vision is rare in vertebrates ❖ Photoreceptors of vertebrates are generally much noisier than those of insects ❖ Monochromatic vision default at night (Warrant and Somanathan, 2022)
  • 54. HOW INSECT FIND SOLUTION TO COLOUR VISION ➢ Discriminating colour in dim light - low visual SNR ➢ Solution lies in optical and neural strategies ➢ Increasing the photon catch or Reducing the noise ➢ Insects have evolved many such strategies o Optically - signal amplitude can be improved by having an eye design that captures more light o Peripheral neural mechanisms that sum photons of light in time and space (Warrant and Somanathan, 2022)
  • 55. SUMMATION SPATIAL SUMMATION ❑ Instead of each channel collecting photons ❑ Activation of specialized laterally spreading neuron - couple visual channels into groups TEMPORAL SUMMATION ❑ Increasing longer exposure time ❑ Responding more slowly and building up a brighter image ❑ Fast-moving object, can be significantly degraded ➢ Summation improve the visual SNR in dim light ➢ Coarser and slower features of the world at the expense of the finer and faster features. ➢ Absence of summation nothing seen (Laughlin and Weckstrom, 1993)
  • 56. A L G O R I T H M I N S P I R E D B Y N O C T U R N A L V I S U A L P R O C E S S I N G ❑ Amplification of primary image signals to optimized spatio– temporal summation to reduce noise ❑ Increases the reliability of video collected in dim light, including the preservation of color (Warrant et al., 2014)
  • 57. HAWK MOTH NOCTURNAL COLOUR VISION ❖ Most hawkmoth are active at dawn and dusk ❖ To find and recognize rewarding flowers independent of the colour of the illuminating light – colour constant visual system ❖ Behavioural experiments shows that the nocturnal hawkmoth Deilephila elpenor uses colour vision to discriminate coloured stimuli at intensities corresponding to dim starlight (0.0001 cd m-2) ❖ First report for nocturnal colour vision in animals (Kelber et al., 2002)
  • 58. NOCTURNAL COLOUR VISION ❖ Generally assumed nocturnal flowers attract insects by their high intensity contrast to the back ground of green leaves ❖ White nocturnal flowers absorb ultraviolet light ❖ Rhodopsins have some sensitivity in the ultraviolet range ❖ Flowers are therefore not maximally bright to an insect eye (Warrant and Somanathan, 2022)
  • 59. E X P E R I M E N T O N CO LO U R V I S I O N ❖ Trained D. elpenor to associate a reward of sugar solution with a colour, either blue or yellow to a human observer ❖ Five different light intensities ranging from mid-dusk (1cdm-2) to dim starlight (0.0001cdm-2) (Kelber et al., 2002)
  • 60. E X P E R I M E N T O N CO LO U R V I S I O N Choice frequencies in tests after training to blue between humans and D. elpenor Choice frequencies in tests after training to yellow between humans and D. elpenor (Kelber et al., 2002)
  • 61. ILLUMINATION AND COLOUR CONSTANCY ❑ Discrimination was tested in two different illumination colours, white and broad-spectrum yellow ❑ Without colour constancy moths would confuse these two stimuli Moths trained to green stimuli (b) and turquoise stimuli (c) chose correctly in tests under two different illuminations, `white' (left) and `yellow' (right) (Kelber et al., 2002)
  • 62. TWO TYPES OF OMMATIDIA ✓ D. elpenor has two different types of ommatidium in the eye: Both types have seven proximal green receptors - one has two distal blue receptors, and other has two distal ultraviolet receptors. ✓ Quantum catches for the ultraviolet photoreceptor were especially low. ✓ 0.0001 cd/m2 ✓ Photon shot noise more ✓ Reliable discrimination of blue from grey is impossible under these conditions - spatial and temporal summation Quantum catches - D. elpenor photoreceptors (Kelber et al., 2002)
  • 63. COLOUR VISION IN BEES ❖ Colour vision in the Apis mellifera - model ❖ Three spectral classes of photoreceptors - ultraviolet (344 nm), blue (436 nm), and green (556 nm) portions of the spectrum enable colour vision ❖ Colour perception reduced in light levels (Vijayan et al., 2023)
  • 64. COLOUR VISION IN ROCK BEE Nocturnal foraging in A. dorsata - lunar phase - third quarter maximum activity ❖ Colour vision thresholds of nocturnally active insects ❖ The violet, blue and green bands represent trichromacy in these species (Vijayan et al., 2023)
  • 65. EXPERIMENT ON COLOUR VISION IN ROCK BEE (Vijayan et al., 2023)
  • 66. MEGALOPTA GENALIS ✓ Sweat bee Megalopta (Hymenoptera: Halictidae) ✓ Nocturnal behaviour is a successful strategy ✓ Megalopta performs visually guided flight with apposition eyes ✓ Forage only when the sun is down: shortly before sunrise, and shortly after sunset (Warrant et al., 2004) ✓ Megalopta nests are often found under thick canopy in the understory dim in bright daylight.
  • 67. ❖ Female Megalopta genalis have relatively large eyes ❖ Corneal facet diameters are 1.8 times larger, and rhabdom diameters 4–5 times larger, than those of diurnal halictids (Greiner et al., 2004) ❖ These optical adaptations together make the eyes of M. genalis 27 times more sensitive than those of their diurnal counterparts MEGALOPTA GENALIS
  • 68. MEGALOPTA VS LASIOGLOSSUM ❖ (a, b) Responses to single photons ❖ Bump amplitude is larger, and the bump time course much slower, in M. genalis than in L. leucozonium ❖ In light-adapted conditions (c, d), both species reach the same maximum contrast gain per unit bandwidth, although L. leucozonium - broader bandwidth ❖ In dark-adapted conditions (e, f), M. genalis has a much higher contrast gain per unit bandwidth photoreceptor responses ❖ Larger bumps of M. genalis - Photoreceptor’s gain of transduction is greater compared with diurnal species Large bumps in nocturnal crane flies (Stavenga et al., 1993), cockroaches (Greiner et al., 2004) ,bees (Matsuura, 1999)
  • 69. NEURAL STRATERGY ❑ Spatial summation, which improves visual reliability by grouping signals from neighbouring photoreceptors ❑ Temporal summation, which does so by increasing integration times ❑ Otherwise plagued by photon noise ❑ Trade resolution for sensitivity is if it improves visual performance (Snyder et al., 1977)
  • 70. LOW LIGHT LEVELS AND MEGALOPTA FLIGHT PERFORMANCE ✓ Bright light landings were always quick ✓ Low light landings included both quick and slow samples ✓ Nocturnal hovering insect's hawkmoths, lose acuity to spatial pooling ✓ Even as light levels and acuity drop, bees cannot slow their approaches ✓ Bees learn the speed at which landmark images move across the retina - goal (Cartwright and Collett, 1979)
  • 71. E X P E R I M E N T O N M E G A L O P TA – L A N D M A R K O R I E N TAT I O N ✓ Orientation flight - Spatial arrangement of landmarks around the nest entrance ✓ Five nests in a row, about 1 m above the forest floor ✓ Nest Leaving light intensity – 0.01 cd/m2 ✓ Returning intensity - 0.0001 cd/m2 (Zeil et al., 1996) Two landmark-manipulation experiment
  • 72. OPTICAL PARAMETER IN MEGALOPTA N=1.13( 𝝅 𝟒 )∆p2 D2 k𝝉 ∆ t‫׬‬ 𝟏 − 𝒆−𝑲𝑹 𝝀 𝒍 /( 𝝀)𝒅𝝀 (Souza et al.,1989) ✓ Megalopta eyes have morphological, optical, and electrophysiological characteristics - suit to nocturnal life ✓ In Megalopta 0.15 photons are absorbed by a single green receptor - one integration time ✓ In Apis at the same intensity, N = 0.0053 photons Acceptance angle - rhabdomere - range of angles from which it can effectively absorb light
  • 73. N O C T R U N A L C A R P E N T E R B E E ✓ Xylocopa tranquebarica nocturnal activity on moonlit nights (Burgett and Sukumalanand, 2000) ✓ Xylocopa tenuiscapa and Xylocopa leucothorax are potential competitors ✓ Exploit that flower for pollen and nectar rewards at night (Hopkins et al., 2000)
  • 74. F L I G H T A C T I V I T Y O F T H R E E S P E C I E S O F C A R P E N T E R B E E S (Somanathan et al., 2007)
  • 75. L A N D M A R K O R I E N TAT I O N E X P E R I M E N T I N X Y L O C O PA (Somanathan et al., 2007)
  • 76. (Somanathan et al., 2007) ANATOMY OF NOCTRUNAL CARPENTER BEE EYES Optical sensitivities - X. tranquebarica has eyes that are 27 times more sensitive than X. leucothorax and 9 times more sensitive than those of X. tenuiscapa OPTICAL SENSITIVITY
  • 77. OCELLI OF THREE CARPENTER BEE X. leucothorax (a, d) X. tenuiscapa (b, e) X. tranquebarica (c, f) Tracheal tapetum in the ocelli of X. tranquebarica others instead have dark pigmentation below the ocellar retina (Somanathan et al., 2009)
  • 78. PATH INTEGRATION ✓ Path integration - Cataglyphis ants ✓ Path integration - leaving a starting point, such as a nest, an animal updates an accumulator that keeps a running tally of its current direction and distance - path back to starting point (Collett and Collett, 2000)
  • 79. N AV I G AT I O N U S I N G C A N O P Y PAT T E R N ❖ Parastrachia japonensis (Heteroptera: Parastrachiidae) ❖ Female leaves her burrow to find drupes - Schoepfia jasminodora (Olacaceae: Rosidae: Santales) ❖ It inserts the proboscis into the drupe and drags - burrow ❖ Direct celestial cues are likely to be unreliable – canopy pattern used Sudden 180◦ rotation of an artificial canopy gap in the roof of its box - 12 bugs (Hironaka et al., 2008)
  • 80. NOCTURNAL BULL ANTS AND ITS NAVIGATION Study revealed that nocturnal bull ants (Myrmecia midas) can navigate using the very faint polarization pattern produced by moonlight (Freas et al., 2024)
  • 81. NOCTURNAL INSECT MIGRATION ➢ Europe’s death’s-head hawkmoths (Acherontia Atropos) ➢ 2,000 miles from Europe to Africa ➢ 14 hawkmoths and recorded their precise GPS locations from a light aircraft - 33.8km/hour ➢ Combination of visual landmarks and Earth’s magnetic field + Favorable tail winds (Menz et al., 2022)
  • 82. NOCTURNAL PEST ➢ Many insects are nocturnal pest, including bed bugs, earwigs, moths, cockroaches etc., ➢ According to research, Fall Armyworm outbreak could potentially lead to maize production losses ranging from 4.1 to 17.7 mil.tons/year 12 maize- producing countries in Sub-Saharan Africa (Rwomushana et al., 2018) ➢ Mosquito-borne diseases cause significant economic damage globally, costing billions of dollars each year
  • 83. LIGHT TRAP ➢ Insects see ultraviolet (UV) radiation ➢ Nocturnal insects are often attracted to light sources that emit large amounts of UV radiation ➢ Devices that exploit this behavior, such as light traps for forecasting pest outbreaks, and electric insect killers – developed ➢ Lamps - yellow illumination effectively to control the activity of nocturnal moths (Shimoda and Honda , 2013)
  • 84. YELLOW FLUORESCENT LAMPS (Shimoda and Honda , 2013) Flower cultivation facilities with yellow fluorescent lamps A: Carnation B: Chrysanthemum
  • 85. LEDS IN PEST CONTROL ❖ Electrical current passes through a microchip - emit light ❖ Different colors - semiconductor materials that emit photons at different wavelengths ❖ Wide spectrum of colours - red, green, and blue (RGB) light ADVANTAGES ✓ Species-specific response ✓ Precise wavelength control ✓ Improved attraction rates ✓ Reduced energy consumption ✓ Heat reduction (Wakefield et al., 2016)
  • 86. NOCTURN AL INSECT MONITORING METHODS (Roy et al., 2024) ❑ Search Light trap ❑ Vertical-looking entomological radar ❑ Camera trap The IMR (Insect Monitoring Radar) Tower, designed by Dr. Christopher M. Kaltenbach, is inspired by the UAE’s historic watchtowers
  • 87. NOCTURN AL INSECT MONITORING METHODS (Roy et al., 2024) AUTOMATED SENSOR ❑ Insects - high diversity - 80% ❑ Sensors comprise a light to attract insects, a camera for collecting images and a computer for scheduling, data storage and processing ❑ Metadata is important - balance the capture - power and data storage limitations
  • 88. NOCTURN AL INSECT MONITORING METHODS (Sun et al., 2022) SEARCH LIGHT TRAP ❑ Searchlight trapping is a method that uses light to attract and trap migratory insects ❑ Three species of rice migratory pests (Cnaphalocrocis medinalis, Sogatella furcifera, and Nilaparvata lugens) cause severe yield and economic losses to rice food every year in china
  • 89. NOCTURN AL INSECT MONITORING METHODS (Roy et al., 2024) Machine learning workflow to analyse moth camera trap data
  • 90. NOCTURN AL INSECT MONITORING METHODS (Cheng et al., 2018; Gao et al., 2024) RADAR ❑ Migration is a key process in the numerous insect species – Pests ❑ Identification of insect migrants is important ❑ Vertical-looking radars (VLRs) - monitor high- altitude insect migration ❑ The VLRs emit a vertically pointing, linear- polarized, narrow-angle conical scan ❑ Insect migrate in tightly clustered – bright moon nights – clouds were sparse
  • 91. IMPACT OF ARTIFICIAL LIGHT AT NIGHT ❖ Localized illumination of nocturnal landscapes by anthropogenic sources of light ❖ Global surveys - 8% above the natural level (Falchi et al., 2016) ❖ Insect diversity and abundance - rapid decline (Hallmann et al., 2017) ❖ In Nocturnal insects - spatial disorientation and attraction ❖ Fireflies ,Click beetles and Glowworms - vulnerable to artificial illumination
  • 92. TEMPORAL DISORIENTATION ❖ ALAN - temporal disorientation, desynchronization of organisms from their biorhythms (Saunders, 2012) ❖ Insect - circadian patterns of activity (foraging, reproduction, migration, etc.) - light cycle ❖ Nocturnal insects - emergence time , feeding and courtship - dictated by internal clocks - ambient light (Saunders, 2009)
  • 93. DESENSITISATION ❖ Nocturnal insects - highly sensitive visual systems – not function well in illuminated environment ❖ Some insects like Myrmecia ants are capable of flexible, rapid light adaptation (Narendra et al., 2013) ❖ Many photons at once, some insects - temporarily dazzled or permanently blinded ❖ Gryllus bimaculatus - exposure to bright UV light structural degeneration of photoreceptors
  • 94. ALAN ON BIOLUMINESCENT INSECTS ✓ 2000 lampyrid species ✓ Adult fireflies - employ bioluminescence - courtship signal ✓ Click beetles (Coleoptera: Elateridae), Railroad worm (Coleoptera: Phengodidae), and Fungus gnat (Diptera: Keroplatidae), employ bioluminescence as an aposematic signal or predatory lure (Rochow, 2007)
  • 95. DORSAL LIGHT RESPONSE  Interaction between flying insects and artificial light – “Drawn like a moth to a flame”  The dorsal-light-response is a behavior where flying insects keep their top side the dorsal side pointed at the brightest area in their vision  Artificial light is an ancient method - Roman Empire - 1 AD Three flight patterns 1.Orbiting: a circular flight around the bulb - body slightly tilted to the light. 2. Stalling: a sharp climb away from the ground as the insect faces away from the light source 3.Inversion: a dive to the ground (Owens and Lewis, 2018)
  • 96. WHY INSECTS DRAWN TO LIGHT ? Insects use the moon as a celestial compass cue to navigate - artificial light sources Thermal radiation from light sources is attractive to flying insects Insects are drawn to light through an escape mechanism -gap in the foliage (Owens and Lewis, 2018)
  • 97. SOLUTIONS TO REDUCE LIGHT POLLUTION ❖ Avoid blue light Insects are particularly attracted to blue wavelengths ❖ Red light It has minimal impact on insects ❖ Shielding: Install fixtures with hoods or baffles to direct light downwards (Schroer et al., 2021)
  • 98. SOLUTIONS TO REDUCE LIGHT POLLUTION ❖ Only light when needed: Turn off lights not in use ❖ Motion sensors: Illuminate when movement detected ❖ Dimmers: Reduce light intensity when necessary ❖ Awareness campaigns: Educate your community about the impacts of light pollution on insects (Schroer et al., 2021)
  • 101. CONCLUSION ❑ Nocturnal insects play the same important role as diurnal insects ❑ They have successfully adapted to the darkest environments ❑ Vision plays a major role ❑ “By reducing light pollution we can save the unseen beauty of night world”