Melophorus bagoti Navigational mechanisms Central place foragers

 Effective, Robust

Wehner and Wehner 1990; Wehner 2003

Navigational mechanisms

 Effective, Robust  Relatively simple

Tiny Brain (1mm3) Parsimonious, Efficient bagoti Gigantiops destructor The ’ navigational toolkit

Path Landmark Systematic Integration navigation Search Path Integration

Food item Direction Distance (celestial compass) (odometer)

Cataglyphis fortis Nest Wehner 1982………………………… 2012 Problems of Path Integration Path - Accumulate errors Integration

Food item Direction Distance (celestial compass) (odometer)

Cataglyphis fortis Nest Problems of Path Integration Path - Accumulate errors Integration - Passive displacement

Food item Direction Distance (celestial compass) (odometer)

Cataglyphis fortis Nest

Wehner and Srinivasan 1981. J. comp. phys Problems of Systematic Path Integration Search - Accumulate errors - Passive displacement

Food item

Cataglyphis fortis Nest

Wehner and Räber 1979. Experencia Cartwright and Collett 1983. J. comp. phys Landmark navigation

Food item

Nest Landmark navigation

Food item Landmarks… Our human way of seeing the world

Nest Our human way of seeing the world

Panorama Landmark Our human way of seeing the world

Distal Panorama Landmark Our human way of seeing the world …..and designing experiments Distal panorama

Proximal Landmarks Our human way of seeing the world …..and designing experiments

Panorama (very flat)

Landmarks Our human way of seeing the w orld

Geometry Features (Extended surfaces) Our human way of seeing the w orld …..and designing experiments

Geometry (Extended surfaces) !

Features ?

A little bit more objective… A little bit more objective… 1 pixel / 4 deg

Melophorus bagoti compound eye: Acuity = 4 deg

(Schwarz et al., 2003. Arthr. Struct & Devlop.) 1 pixel / 4 deg

1 pixel / 4 deg

UV 1 pixel / 4 deg

Green A little bit more objective… A little bit more objective…

Do segregate landmarks and panorama? (Wystrach and Graham 2012) Quantify the modification of the scenery…

2 days

Training

N

10m

F F (Wystrach et al., 2011) Quantify the modification of the scenery…

Reference pictures Compared pictures Map of mismatch (training condition) (Test conditions)

N N

10m Comparisons

F F Relate image difference / behaviour…

Cues widespread on their panoramic view

U turns

Degradation of the matching Zero vector ants (Wystrach et al., 2011) Information available to the

Cues widespread on their panoramic view

Landmark navigation Information available to the animal

Cues widespread on their panoramic view

Visual scene navigation Identify the problem in natural environment Visual scene navigation

Cataglyphis velox

Melophorus bagoti Route following

Cataglyphis velox

Melophorus bagoti

Mangan and Webb 2012. Behav. Ecol. Kohler and Wehner 2005, Neurobiol. of Learn. and Mem. Wystrach, Philippides and Graham (in prep). Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest

Current view Current view

Visual scene navigation Modelling: Zeil et al. 2003. J. Opt. Soc. Am. A; Graham et al. 2010. Cur. Biol. Ant experiment: Wystrach et al. 2011. J. Exp. Psych.

Memory

Nest Best match! Current view Current view

Simply move in the most familiar direction

Visual scene navigation Route following model (Baddeley et al., 2012)

Route following No positional knowledge Simply move in the most familiar direction Homing from novel locations

Melophorus bagoti N

Melophorus bagoti Wystrach et al. 2012. J. Exp. Biol Homing from novel locations

(Cartwright and Collett, 1983; Zeil et al., 2003) Homing from novel locations

(Cartwright and Collett, 1983; Zeil et al., 2003) Homing from novel locations

Simply move as to increase the familiarity

(Cartwright and Collett, 1983; Zeil et al., 2003) Visual scene Feeder navigation

Route following No positional knowledge Simply going in the most familiar direction

Familiar terrain

Nest Homing from novel locations No positional knowledge Simply moving so as the familiarity increase The ants’ navigational toolkit

Path Visual scene Systematic Integration navigation Search

Quite simple …

How can it be so robust ? How can it be so robust ?

Kohler and Wehner 2005. Neurobiol. Learn. Mem. Melophorus bagoti

How can it be so robust ?

Nest

Melophorus bagoti Wystrach et al. 2011. JCPA How can it be so robust ?

Wystrach et al. 2011. JCPA How can it be so robust ?

Nest

Melophorus bagoti Wystrach et al. 2011. JCPA How can it be so robust ? Sensory level: using multiple cues How can it be so robust ? Sensory level: using multiple cues

Path Integration

Direction Distance (celestial compass) (odometer) How can it be so robust ? Sensory level: using multiple cues

Path Integration

Direction Distance

Polarized light Proprioception Sun position (Stride, Ascent Chromatic cues and Descent) Light gradient Optic flow Wind direction

Wehner 2008. Myrmec. news Wittlinger et al., 2006. Science Wystrach and Schwarz, in prep Ronacher 2008. Myrmec. News How can it be so robust ? Sensory level: using multiple cues Visual scene navigation

Skyline Centre of mass Edges orientation Optic flow pattern

Wystrach et al., 2011. Front. in Zool. How can it be so robust ? Sensory level: using multiple cues Visual scene navigation

Panoramic views Wystrach et al., 2011. Front. in Zool. Motor routines Knaden et al. 2006. Curr. biol Odor cues Steck et al., 2009. Front. in Zool. Magnetic cue Buehlmann et al., 2012. Plos one

Vibratory cue Cataglyphis noda Buehlmann et al., 2012. Plos one credit: Max Planck Institute for Chemical Ecology/Badeke

How can it be so robust ? Sensory level: using multiple cues Learnt information

Panoramic views Wystrach et al., 2011. Front. in Zool. Motor routines Knaden et al. 2006. Curr. biol Odor cues Steck et al., 2009. Front. in Zool. Magnetic cue Buehlmann et al., 2012. Plos one

Vibratory cue Cataglyphis noda Buehlmann et al., 2012. Plos one credit: Max Planck Institute for Chemical Ecology/Badeke

How can it be so robust ? Sensory level: using multiple cues

Chromatic gradient Panoramic views Proprioception Polarized light Vibratory cue Light gradient Optic flow Odor cues Sun position Motor routines Wind direction Magnetic cues

Path Learnt Systematic Integration information Search

Emergency plan! How can it be so robust ? Output level: combining strategies

Path Learnt Systematic Integration information Search ?

Behavior How can it be so robust ? Output level: combining strategies

Path Learnt Systematic Integration information Search

Map-like representation?

Behavior How can it be so robust ? Output level: combining strategies

Path Learnt Systematic Integration information Search

Learnt views operate independently from PI Map-like representation? (Kohler and Wehner, 2005) PI operate independently from views (Andel and Wehner, 2002)

Behavior N F

X 10 How can it be so robust ? Output level: combining strategies

Path Learnt Systematic Integration information Search

Learnt views operate independently from PI Map-like representation? (Kohler and Wehner, 2005) PI operate independently from views (Andel and Wehner, 2002)

N F How can it be so robust ? Output level: combining strategies

Path Learnt Systematic Integration information Search

How do the different Behavior systems interact?

Wehner 2008, Myrmec. News. How can it be so robust ? Output level: combining strategies

PI Views

Zero vector Nest

Melophorus bagoti Wystrach and Legge, in prep How can it be so robust ? Output level: combining strategies

Zero vector Nest

Full vector

Feeder PI Views

Melophorus bagoti Wystrach and Legge, in prep How can it be so robust ? Output level: combining strategies

PI Views Zero vector Nest

Full vector

Melophorus bagoti Wystrach and Legge, in prep How can it be so robust ? Output level: combining strategies

Zero vector Nest

Full vector

Feeder

PI Views Wystrach and Legge, in prep See also Collett 2012. Curr. biol. How can it be so robust ? Output level: combining strategies

PI Views Zero vector Nest

Full vector

Wystrach and Legge, in prep How can it be so robust ? Output level: combining strategies

Zero vector Nest

Full vector

Feeder

PI Views Wystrach and Legge, in prep How can it be so robust ? Output level: combiningweighting strategies

Zero vector Nest

Full vector

Feeder

Wystrach and Legge, in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Current Familiarity

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Current Familiarity

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity continuous

How are the Behavior strategies weighted?

Wystrach et al. in prep Weighting the different systems

Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity continuous

How are the Behavior strategies weighted?

Wystrach et al. in prep Path Integration

Feeder Vector length

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest unfamiliar Path Integration terrain Feeder Vector length

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest Path Integration

Feeder Vector length

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest Path Integration

Feeder Path Vector length integration

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest Totally Path Integration unfamiliar Feeder Path Vector length integration terrain ? Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest Path Path Path Integration integration integration Feeder Vector length

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous

Nest Systematic Systematic search search What ants do when they are lost ?

Feeder

Nest

Melophorus bagoti

Wystrach et al. in prep What ants do when they are lost ? Distant release points: unfamiliar environment

Feeder

Zero vector ?

Nest Path Visual scene Systematic Integration navigation Search

Vector length Current Familiarity continuous

Behavior Melophorus bagoti

Wystrach et al. in prep What ants do when they are lost ? Distant release points: unfamiliar environment

Feeder

Zero vector

Nest

Melophorus bagoti

Wystrach et al. in prep Distant release points: unfamiliar environment

Zero vector

Nest Zero vector

Feeder

Wystrach et al. in prep Distant release points: unfamiliar environment

Backtracking

Zero vector

Nest Zero vector

Zero-vector ants backtrack towards the feeder Feeder

Wystrach et al. in prep When do ants backtrack ?

Path Backtracking Visual scene Systematic Integration Navigation Search Current Vector length continuous ? Familiarity

Output When do ants backtrack ?

Feeder Feeder

Zero vector Residual PI Nest Nest ants vector

- On unfamiliar terrain… - When PI vector is small…

Wystrach et al. in prep When do ants backtrack ?

Recent view of the nest vicinity!

Feeder Feeder Feeder

Zero vector Zero vector Zero vector Nest Nest Nest ants ants ants

- On unfamiliar terrain… - When PI vector is small… - Recent view of the nest vicinity Wystrach et al. in prep When do ants backtrack ?

Path Backtracking Visual scene Systematic Integration Navigation Search Recent view of Current Vector length the nest vicinity continuous Familiarity

Output - On unfamiliar terrain… - When PI vector is small… - Recent view of the nest vicinity Robust !

Path Path Path Integration integration integration Feeder Vector length

Visual scene navigation Current Familiarity

Systematic Familiar Search terrain continuous Systematic Systematic search search Backtracking

Nest Recent view of the nest vicinity Backtracking Backtracking Robust !

Path Path integration integration Feeder Complex Decision making ?

A distributed system with simple interactions

Path scene Systematic Integration navigation Search Familiar terrain Systematic Systematic search search

No need of a higher Nest representation

Backtracking Backtracking Simple solutions behind apparently complex behaviour

Exploit complex visual information without landmark recognition

Follow complex routes without positional knowledge

Home from novel locations without cognitive map

Path scene Systematic Complex decision making Integration navigation Search without higher representations CONCLUSION Top-Down Human way of seeing the world

(Landmarks) (Cognitive map)

Literature What is their ecological problem ? Hypotheses How do they solve it? use landmarks! Behavioural criterion

Animal’s Umwelt

Result Bottom-up Insects use landmarks! Chittka et al., 2012 Wystrach and Graham, 2012

Sebastian Schwarz Thanks Patrick Schultheiss

Alice Baniel Ken Cheng

Paul Graham The ants you Michael Mangan

Bart Baddeley Andy Philippides