Excerpt

Understanding the organization of the vertebrate retina has been the goal of many talented visual scientists during the past 100 years. With Cajal's (1892) anatomic descriptions of the cell types that constitute the retina in a number of vertebrate species, and with an early understanding of the role of visual purple in photochemistry in combination with psychophysical studies of adaptation and color vision, we had in the sixties the rudiments of an understanding of how the retina might be organized and functioning.

To go further, though, we were beginning to need detailed information of neural circuits that underlie these functions. It was the advent of electron microscopy, microelectrode recording techniques, and pharmacology that then allowed us an era of very rapid advancement. The purpose of this Electronic Tutorial is to summarize these recent advances and to describe our present understanding, based primarily on anatomic investigations, of the Neural Organization of the Mammalian Retina. As time goes on we have been inviting other authors to write chapters on their speciality concerning the retina or other visual pathways. A great addition has been a section on Psychophysics of Vision which we hope will be of general information to all interested in learning the basics of visual perception.

Contents

• About the Editors
  Helga Kolb, Eduardo Fernandez, Bryan Jones, and Ralph Nelson.
  Last Update: October 8, 2024.
• Acknowledgement
  Helga Kolb, Eduardo Fernandez, Bryan Jones, and Ralph Nelson.
  Last Update: April 28, 2026.
• Part I: Foundations
  • Introduction
    Helga Kolb.
  • Gross Anatomy of the Eye
    Helga E. Kolb.
    Created: May 1, 2005; Last Update: October 10, 2024.
  • Simple Anatomy of the Retina
    Helga Kolb.
    Created: May 1, 2005; Last Update: January 31, 2012.
  • How the Retina Works^*
    Helga E. Kolb and Ralph F. Nelson.
    Created: February 1, 2003; Last Update: July 4, 2025.
• Part II: Anatomy and Physiology of the retina
  • The Retinal Pigment Epithelium
    Olaf Strauss.
    Created: January 26, 2011.
  • Photoreceptors
    Ralph Nelson and Helga E. Kolb.
    Created: February 1, 2003; Last Update: April 22, 2026.
  • OUTER PLEXIFORM LAYER
    Helga Kolb.
    Created: May 1, 2005; Last Update: October 8, 2011.
  • Inner Plexiform Layer
    Helga Kolb.
    Last Update: March 25, 2012.
  • Morphology and Circuitry of Ganglion Cells by Helga Kolb
    Helga Kolb.
  • Visual Responses of Ganglion Cells
    Ralph Nelson.
    Created: May 1, 2001; Last Update: April 10, 2007.
  • Melanopsin-expressing, Intrinsically Photosensitive Retinal Ganglion Cells (ipRGCs)
    Dustin M. Graham and Kwoon Y. Wong.
    Created: August 1, 2008; Last Update: November 2, 2016.
  • Glial cells of the Retina by Helga Kolb
    Helga Kolb.
  • The Architecture of the Human Fovea
    Helga Kolb, Ralph F. Nelson, Peter K. Ahnelt, Isabel Ortuño-Lizarán, and Nicolas Cuenca.
    Created: February 7, 2020; Last Update: May 20, 2020.
• Part III: Retinal Circuits
  • Circuitry for Rod Signals through the Retina
    Helga Kolb.
    Created: May 1, 2001; Last Update: September 1, 2009.
  • Cone Pathways through the Retina by Helga Kolb
    Helga Kolb.
  • Roles of Amacrine Cells by Helga Kolb
    Helga Kolb.
  • The Anatomy and Physiology of Direction-Selective Retinal Ganglion Cells
    Jinyue Liu.
    Created: August 20, 2015.
  • AII Amacrine Cells by Mahnoosh Farsaii and Victoria P. Connaughton
    Mahnoosh Farsaii and Victoria P. Connaughton.
  • Midget pathways of the primate retina underlie resolution and red green color opponency
    Helga Kolb.
    Created: May 24, 2007; Last Update: April 4, 2012.
  • S-Cone Pathways by Helga Kolb
    Helga Kolb.
  • Feedback Loops in the Retina
    Helga Kolb.
    Created: May 1, 2005; Last Update: May 4, 2007.
  • Myriad roles for gap junctions in retinal circuits
    Stuart Trenholm and Gautam B. Awatramani.
    Created: September 20, 2019.
• Part IV: Neurotransmitters in the Retina
  • Neurotransmitters in the Retina by Helga Kolb
    Helga Kolb.
  • Glycine Receptor Diversity in the Mammalian Retina
    Silke Haverkamp.
    Created: March 27, 2012.
  • Role of Dopamine in Retinal Function
    Elka Popova.
    Created: May 28, 2020.
• Part V: Phototransduction and Photoreceptor Synaptic Pathways
  • Phototransduction in Rods and Cones
    Yingbin Fu.
    Created: April 1, 2010; Last Update: July 30, 2018.
  • Glutamate and Glutamate Receptors in the Vertebrate Retina
    Victoria Connaughton.
    Created: May 1, 2005; Last Update: May 7, 2007.
  • Bipolar Cell Pathways in the Vertebrate Retina
    Ralph Nelson and Victoria Connaughton.
    Created: May 24, 2007; Last Update: January 20, 2012.
  • GABA_C Receptors in the Vertebrate Retina
    Haohua Qian.
    Created: May 1, 2005; Last Update: June 1, 2007.
  • S-Potentials and Horizontal Cells
    Ido Perlman, Helga Kolb, and Ralph Nelson.
    Created: May 1, 2007; Last Update: December 30, 2011.
• Part VI: Retinal Neurogenesis: Early stages in the development of neurons and pathways: Early stages in the Development of Neurons and Pathways
  • Development of cell types and synaptic connections in the retina by Josh Morgan and Rachel Wong
    Josh Morgan and Rachel Wong.
    Last Update: July 30, 2018.
  • Formation of Early Retinal Circuits in the Inner-Plexiform Layer
    Kevin Ford and Marla Feller.
    Created: May 1, 2005; Last Update: January 16, 2012.
  • Development of Retinal Ganglion Cell Dendritic Structure and Synaptic Connections
    Ning Tian.
    Created: June 13, 2012.
• Part VII: Color Vision
  • Color Vision
    Peter Gouras.
    Created: May 1, 2005; Last Update: July 1, 2009.
• Part VIII: Psychophysics of Vision
  • Psychophysics of Vision by Michael Kalloniatis and Charles Luu
    Michael Kalloniatis and Charles Luu.
    Last Update: August 19, 2011.
  • Visual Acuity
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: June 5, 2007.
  • Temporal Resolution
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: June 5, 2007.
  • Light and Dark Adaptation
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: July 9, 2007.
  • The Perception of Color
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: July 9, 2007.
  • The Perception of Space
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: June 6, 2007.
  • The Perception of Depth
    Michael Kalloniatis and Charles Luu.
    Created: May 1, 2005; Last Update: June 6, 2007.
• Part IX: Brain Visual Areas
  • The Primary Visual Cortex
    Matthew Schmolesky.
    Created: May 1, 2005; Last Update: June 14, 2007.
• Part X: Repair and Regeneration in the visual sytem
  • Regeneration in the Goldfish Visual System
    Sam Nona.
    Created: May 1, 2005; Last Update: June 20, 2007.
  • Regeneration in the Visual System of Adult Mammals
    Yves Sauve and Frederic Gaillard.
    Created: May 1, 2005; Last Update: June 21, 2007.
  • Fetal Tissue Allografts in the Central Visual System of Rodents
    Frederic Gaillard and Yves Sauve.
    Created: May 1, 2005; Last Update: June 25, 2007.
• Part XI: Electrophysiology
  • VISUALLY EVOKED POTENTIALS
    Donnell J. Creel.
    Created: March 1, 2012; Last Update: July 14, 2015.
  • The Electroretinogram: ERG
    Ido Perlman.
    Created: May 1, 2001; Last Update: June 27, 2007.
  • The Electroretinogram and Electro-oculogram: Clinical Applications
    Donnell J. Creel.
    Last Update: July 14, 2015.
  • Visual and Auditory Anomalies Associated with Albinism
    Donnell J. Creel.
    Created: June 18, 2015.
• Part XII: Investigations of Human Retinal Disease
  • Retinal Degeneration, Remodeling and Plasticity
    Bryan William Jones, Robert E. Marc, and Rebecca L. Pfeiffer.
    Created: October 28, 2016.
  • Cellular Remodeling in Mammalian Retina Induced by Retinal Detachment
    Steven K. Fisher, Geoffrey P. Lewis, Kenneth A. Linberg, Edward Barawid, and Mark R. Verardo.
    Created: May 1, 2005; Last Update: July 3, 2007.
  • Age-Related Macular Degeneration (AMD)
    Gregory S. Hageman, Karen Gehrs, Lincoln V. Johnson, and Don Anderson.
    Created: January 1, 2008.
  • Diabetic retinopathy and a novel treatment based on the biophysics of rod photoreceptors and dark adaptation
    Geoffrey B. Arden and David J. Ramsey.
    Created: July 14, 2015.
  • What is glaucoma?
    David Križaj.
    Created: May 30, 2019.
• Part XIII: Facts and Figures concerning the human retina by Helga Kolb
  • Facts and Figures Concerning the Human Retina
    Helga Kolb.
    Created: May 1, 2005; Last Update: July 5, 2007.
• Part XIV: Evolution of Phototransduction, Vertebrate Photoreceptors and Retina by Trevor Lamb
  • Evolution of Phototransduction, Vertebrate Photoreceptors and Retina
    Trevor D Lamb.
    Created: May 14, 2013; Last Update: July 10, 2013.
• Part XV: Visual Prostheses
  • Introduction to Visual Prostheses
    Eduardo Fernandez and Richard Normann.
    Created: March 9, 2016.
  • Retinal Prostheses
    Ethan D. Cohen.
    Created: March 19, 2018.
• Part XVI: Anterior Segment
  • Crystalline Lens and Cataract
    Joah F. Aliancy and Nick Mamalis.
    Created: August 15, 2017.
  • Intraocular Lenses for Cataract Surgery
    Jason Nguyen and Liliama Werner.
    Created: August 5, 2017.
• Part XVII: Refractive Errors
  • The Science Behind Myopia
    Brittany J. Carr and William K. Stell.
    Created: November 7, 2017.