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Peter Sterling

University of Pennsylvania, Neuroscience, Faculty Member

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My broad goal has been to learn how the brain is designed – to understand its functional architecture. My research has spanned the full range of scales, from nanoscopic (synaptic vesicles), to microscopic (neural circuits), to macroscopic (regional neuroanatomy and behavior). My laboratory focused on retinal structure and function (Sterling, 2013), but my theoretical interests extended to basic issues of physiological regulation and behavior, leading to the concept of allostasis (Sterling, 2004, 2011). The two lines are now integrated in a book written with Prof. Simon Laughlin, Principles of Neural Design (2015). Investigations continue in a Massachusetts garden and on a farm in western Panamá.
Selected Publications
Sterling P, Laughlin, Simon. : Principles of Neural Design. MIT Press. May 2015.

Sterling P: Homeostasis vs. Allostasis: Implications for brain function and mental disorders. JAMA Psychiatry 71(10): 1192-3, 2014.

Sterling P: Some Principles of Retinal Design: The Proctor Lecture. Invest. Ophthalmol. Vis. Sci. 54(3): 2267-2275, 2013.

Perge J, Niven J, Mugnaini E, Balasubramanian V, Sterling P: Why do axons vary in caliber? J. Neurosci. 32(2): 626-38, 2012.

Sterling P: Allostasis: A model of predictive regulation. Physiol. Behav. 106(1): 5-15, 2012.

Tkačik G, Garrigan P, Ratliff C, Milčinski G, Klein JM, Seyfarth LH, Sterling P, Brainard DH, Balasubramanian V: Natural images from the birthplace of the human eye. PLoS One 6(6): e20409, 2011.

Ratliff CP, Borghuis BG, Kao YH, Sterling P, Balasubramanian V: Retina is structured to process an excess of darkness in natural scenes. Proc Natl Acad Sci U S A 107: 17368-73, 2010.

Borghuis BG, Sterling P, Smith RG: Loss of sensitivity in an analog neural circuit. J. Neurosci. 29(10): 3045-3058, 2009.

Perge J, Koch K, Miller R, Sterling P, Balasubramanian V: How the optic nerve allocates space, energy, capacity, and information. J. Neurosci. 29(24): 7917-7928, 2009.

Borghuis BG, Ratliff CP, Smith RG, Sterling P, Balasubramanian V: Design of a neuronal array. J. Neurosci. 28(12): 3178-3189, 2008.

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Papers by Peter Sterling

Principles of Neural Design

How Bigger Brains Are Organized

The MIT Press eBooks, Jun 30, 2015

Beyond the Retina: Pathways to Perception and Action

The MIT Press eBooks, Jun 30, 2015

Anatomical organization of the brachial spinal cord of the cat. I. The distribution of dorsal root fibers

Brain Research, Feb 1, 1967

The present study has attempted to correlate the distribution and orientation of the motoneuron d... more The present study has attempted to correlate the distribution and orientation of the motoneuron dendrites with the somatotopically organized distribution of their cell bodies. Despite local variations, the predominant dendritic orientation within the motoneuronal cell groups is longitudinal. Retrograde changes after section of various combinations of peripheral nerves show that motoneurons supplying the intrinsic flexor muscles of the forelimb are aligned longitudinally in the dorsolateral regions of the lateral motoneuronal cell groups from C6-T1. Extensor motoneurons are similarly aligned with each other but lie ventral to the flexor motoneurons. Motoneurons supplying the girdle musculature lie still more ventrally and medially. The cell bodies and dendrites intermingle, forming a longitudinal plexus which is supplied by afferents whose terminal arborizations also run longitudinally. The possible roles of these arrangements were discussed with reference to both the maintenance of specificity of the afferent supply to the motoneurons and the activation of synergistic motoneurons.

Why a Bigger Brain?

The MIT Press eBooks, Jun 30, 2015

What Engineers Know about Design

The MIT Press eBooks, Jun 30, 2015

How Photoreceptors Optimize the Capture of Visual Information

The MIT Press eBooks, Jun 30, 2015

Information Processing in Protein Circuits

The MIT Press eBooks, Jun 30, 2015

Current Biology, Feb 1, 2021

Interview with Peter Sterling, author of Principles of Neural Design and What Is Health?

What Is Health?

Anatomy and physiology of goldfish oculomotor system. I. Structure of abducens nucleus

Journal of Neurophysiology, May 1, 1977

Receptive fields and functional architecture in the retina

The Journal of Physiology, Jun 12, 2009

Functional architecture of the striate cortex is known mostly at the tissue level – how neurons o... more Functional architecture of the striate cortex is known mostly at the tissue level – how neurons of different function distribute across its depth and surface on a scale of millimetres. But explanations for its design – why it is just so – need to be addressed at the synaptic level, a much finer scale where the basic description is still lacking. Functional architecture of the retina is known from the scale of millimetres down to nanometres, so we have sought explanations for various aspects of its design. Here we review several aspects of the retina's functional architecture and find that all seem governed by a single principle: represent the most information for the least cost in space and energy. Specifically: (i) why are OFF ganglion cells more numerous than ON cells? Because natural scenes contain more negative than positive contrasts, and the retina matches its neural resources to represent them equally well; (ii) why do ganglion cells of a given type overlap their dendrites to achieve 3-fold coverage? Because this maximizes total information represented by the array – balancing signal-to-noise improvement against increased redundancy; (iii) why do ganglion cells form multiple arrays? Because this allows most information to be sent at lower rates, decreasing the space and energy costs for sending a given amount of information. This broad principle, operating at higher levels, probably contributes to the brain's immense computational efficiency.

What Is Health?: Allostasis and the Evolution of Human Design

Patchy mosaic of L and M cones permits efficient tradeoff between spatial and color vision

OSA Annual Meeting, 1993

Optimal sampling for the luminance channel requires a fine array of cones with identical waveleng... more Optimal sampling for the luminance channel requires a fine array of cones with identical wavelength sensitivity. Trichromacy creates a problem because at each transition between M and L cones their different wavelength sensitivities must either introduce a spurious edge or else distort the magnitude of an existing edge.

Why I joined the Freedom Rides

Current Biology, Jun 1, 2021

Peter Sterling expands upon his recent Q & A article by discussing his participation in the F... more

Micro circuitry of the beta x ganglion cell in dark adaptation

Neighboring Retinal Ganglion Cells Provide Independent Channels for Contrast Detection

Investigative Ophthalmology & Visual Science, May 1, 2005

Responses of Different Retinal Ganglion Cell Types to Motion of 'Natural' Scenes

Investigative Ophthalmology & Visual Science, 2005

Structure and function of neural networks in the retina

Proceedings, annual meeting, Electron Microscopy Society of America, 1988

The synaptic connections in cat retina that link photoreceptors to ganglion cells have been analy... more The synaptic connections in cat retina that link photoreceptors to ganglion cells have been analyzed quantitatively. Our approach has been to prepare serial, ultrathin sections and photograph en montage at low magnification (˜2000X) in the electron microscope. Six series, 100-300 sections long, have been prepared over the last decade. They derive from different cats but always from the same region of retina, about one degree from the center of the visual axis. The material has been analyzed by reconstructing adjacent neurons in each array and then identifying systematically the synaptic connections between arrays. Most reconstructions were done manually by tracing the outlines of processes in successive sections onto acetate sheets aligned on a cartoonist's jig. The tracings were then digitized, stacked by computer, and printed with the hidden lines removed. The results have provided rather than the usual one-dimensional account of pathways, a three-dimensional account of circui...

Author response: Predictive regulation and human design

Principles of Neural Design

How Bigger Brains Are Organized

The MIT Press eBooks, Jun 30, 2015

Beyond the Retina: Pathways to Perception and Action

The MIT Press eBooks, Jun 30, 2015

Anatomical organization of the brachial spinal cord of the cat. I. The distribution of dorsal root fibers

Brain Research, Feb 1, 1967

The present study has attempted to correlate the distribution and orientation of the motoneuron d... more The present study has attempted to correlate the distribution and orientation of the motoneuron dendrites with the somatotopically organized distribution of their cell bodies. Despite local variations, the predominant dendritic orientation within the motoneuronal cell groups is longitudinal. Retrograde changes after section of various combinations of peripheral nerves show that motoneurons supplying the intrinsic flexor muscles of the forelimb are aligned longitudinally in the dorsolateral regions of the lateral motoneuronal cell groups from C6-T1. Extensor motoneurons are similarly aligned with each other but lie ventral to the flexor motoneurons. Motoneurons supplying the girdle musculature lie still more ventrally and medially. The cell bodies and dendrites intermingle, forming a longitudinal plexus which is supplied by afferents whose terminal arborizations also run longitudinally. The possible roles of these arrangements were discussed with reference to both the maintenance of specificity of the afferent supply to the motoneurons and the activation of synergistic motoneurons.

Why a Bigger Brain?

The MIT Press eBooks, Jun 30, 2015

What Engineers Know about Design

The MIT Press eBooks, Jun 30, 2015

How Photoreceptors Optimize the Capture of Visual Information

The MIT Press eBooks, Jun 30, 2015

Information Processing in Protein Circuits

The MIT Press eBooks, Jun 30, 2015

Current Biology, Feb 1, 2021

Interview with Peter Sterling, author of Principles of Neural Design and What Is Health?

What Is Health?

Anatomy and physiology of goldfish oculomotor system. I. Structure of abducens nucleus

Journal of Neurophysiology, May 1, 1977

Receptive fields and functional architecture in the retina

The Journal of Physiology, Jun 12, 2009

Functional architecture of the striate cortex is known mostly at the tissue level – how neurons o... more Functional architecture of the striate cortex is known mostly at the tissue level – how neurons of different function distribute across its depth and surface on a scale of millimetres. But explanations for its design – why it is just so – need to be addressed at the synaptic level, a much finer scale where the basic description is still lacking. Functional architecture of the retina is known from the scale of millimetres down to nanometres, so we have sought explanations for various aspects of its design. Here we review several aspects of the retina's functional architecture and find that all seem governed by a single principle: represent the most information for the least cost in space and energy. Specifically: (i) why are OFF ganglion cells more numerous than ON cells? Because natural scenes contain more negative than positive contrasts, and the retina matches its neural resources to represent them equally well; (ii) why do ganglion cells of a given type overlap their dendrites to achieve 3-fold coverage? Because this maximizes total information represented by the array – balancing signal-to-noise improvement against increased redundancy; (iii) why do ganglion cells form multiple arrays? Because this allows most information to be sent at lower rates, decreasing the space and energy costs for sending a given amount of information. This broad principle, operating at higher levels, probably contributes to the brain's immense computational efficiency.

What Is Health?: Allostasis and the Evolution of Human Design

Patchy mosaic of L and M cones permits efficient tradeoff between spatial and color vision

OSA Annual Meeting, 1993

Optimal sampling for the luminance channel requires a fine array of cones with identical waveleng... more Optimal sampling for the luminance channel requires a fine array of cones with identical wavelength sensitivity. Trichromacy creates a problem because at each transition between M and L cones their different wavelength sensitivities must either introduce a spurious edge or else distort the magnitude of an existing edge.

Why I joined the Freedom Rides

Current Biology, Jun 1, 2021

Peter Sterling expands upon his recent Q & A article by discussing his participation in the F... more

Micro circuitry of the beta x ganglion cell in dark adaptation

Neighboring Retinal Ganglion Cells Provide Independent Channels for Contrast Detection

Investigative Ophthalmology & Visual Science, May 1, 2005

Responses of Different Retinal Ganglion Cell Types to Motion of 'Natural' Scenes

Investigative Ophthalmology & Visual Science, 2005

Structure and function of neural networks in the retina

Proceedings, annual meeting, Electron Microscopy Society of America, 1988

The synaptic connections in cat retina that link photoreceptors to ganglion cells have been analy... more The synaptic connections in cat retina that link photoreceptors to ganglion cells have been analyzed quantitatively. Our approach has been to prepare serial, ultrathin sections and photograph en montage at low magnification (˜2000X) in the electron microscope. Six series, 100-300 sections long, have been prepared over the last decade. They derive from different cats but always from the same region of retina, about one degree from the center of the visual axis. The material has been analyzed by reconstructing adjacent neurons in each array and then identifying systematically the synaptic connections between arrays. Most reconstructions were done manually by tracing the outlines of processes in successive sections onto acetate sheets aligned on a cartoonist's jig. The tracings were then digitized, stacked by computer, and printed with the hidden lines removed. The results have provided rather than the usual one-dimensional account of pathways, a three-dimensional account of circui...

Author response: Predictive regulation and human design