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Fish Do Not Feel Pain and Its Implications for Understanding Phenomenal Consciousness
Biol Philos (2015) 30:149–165 DOI 10.1007/s10539-014-9469-4 Fish do not feel pain and its implications for understanding phenomenal consciousness Brian Key Received: 14 April 2014 / Accepted: 6 December 2014 / Published online: 16 December 2014 Ó The Author(s) 2014. This article is published with open access at Springerlink.com Abstract Phenomenal consciousness or the subjective experience of feeling sen- sory stimuli is fundamental to human existence. Because of the ubiquity of their subjective experiences, humans seem to readily accept the anthropomorphic extension of these mental states to other animals. Humans will typically extrapolate feelings of pain to animals if they respond physiologically and behaviourally to noxious stimuli. The alternative view that fish instead respond to noxious stimuli reflexly and with a limited behavioural repertoire is defended within the context of our current understanding of the neuroanatomy and neurophysiology of mental states. Consequently, a set of fundamental properties of neural tissue necessary for feeling pain or experiencing affective states in vertebrates is proposed. While mammals and birds possess the prerequisite neural architecture for phenomenal consciousness, it is concluded that fish lack these essential characteristics and hence do not feel pain. Keywords Fish Á Pain Á Phenomenal consciousness Á Affective states Á Avoidance learning Á Neocortex Á Pallium Introduction There is a belief in some scientific and lay communities that because fish respond behaviourally to noxious stimuli, then ipso facto, fish feel pain. Sneddon (2011) clearly articulates the logic by stating: ‘‘to explore the possibility of pain perception in nonhumans we use indirect measures similar to those used for human infants who cannot convey whether they are in pain. -
Abnormalities of Grey and White Matter [11C]Flumazenil Binding In
Brain (2002), 125, 2257±2271 Abnormalities of grey and white matter [11C]¯umazenil binding in temporal lobe epilepsy with normal MRI A. Hammers,1,2,3 M. J. Koepp,1,2,3 R. Hurlemann,2 M. Thom,2 M. P. Richardson,1,2,3 D. J. Brooks1 and J. S. Duncan1,2,3 1MRC Clinical Sciences Centre and Division of Correspondence to: Professor John S. Duncan, MA, DM, Neuroscience, Faculty of Medicine, Imperial College, FRCP, National Society for Epilepsy and Institute of 2Department of Clinical and Experimental Epilepsy, Neurology, 33 Queen Square, London WC1N 3BG, UK Institute of Neurology, University College London, London E-mail: [email protected] and 3National Society for Epilepsy MRI Unit, Chalfont St Peter, UK Summary In 20% of potential surgical candidates with refrac- the 16 patients with abnormalities, ®ndings were con- tory epilepsy, current optimal MRI does not identify cordant with EEG and clinical data, enabling further the cause. GABA is the principal inhibitory neuro- presurgical evaluation. Group ®ndings were: (i) transmitter in the brain, and GABAA receptors are decreased FMZ-Vd in the ipsilateral (Z = 3.01) and expressed by most neurones. [11C]Flumazenil (FMZ) contralateral (Z = 2.56) hippocampus; (ii) increased PET images the majority of GABAA receptor sub- FMZ-Vd in the ipsilateral (Z = 3.71) and contralat- types. We investigated abnormalities of FMZ binding eral TLWM (two clusters, Z = 3.11 and 2.79); and in grey and white matter in 18 patients with refrac- (iii) increased FMZ-Vd in the ipsilateral frontal lobe tory temporal lobe epilepsy (TLE) and normal quan- white matter between the superior and medial frontal titative MRI. -
Corticostriatal Interactions During Learning, Memory Processing, and Decision Making
The Journal of Neuroscience, October 14, 2009 • 29(41):12831–12838 • 12831 Mini-Symposium Corticostriatal Interactions during Learning, Memory Processing, and Decision Making Cyriel M. A. Pennartz,1 Joshua D. Berke,2,3 Ann M. Graybiel,4,5 Rutsuko Ito,6 Carien S. Lansink,1 Matthijs van der Meer,7 A. David Redish,7 Kyle S. Smith,4,5 and Pieter Voorn8 1University of Amsterdam, Swammerdam Institute for Life Sciences Center for Neuroscience, 1098 XH Amsterdam, The Netherlands, 2Department of Psychology and 3Neuroscience Program, University of Michigan, Ann Arbor, Michigan 48109, 4Department of Brain and Cognitive Sciences and 5McGovern Institute for Brain Research, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, 6Department of Experimental Psychology, University of Oxford, Oxford OX1 3UD, United Kingdom, 7Department of Neuroscience, University of Minnesota, Minneapolis, Minnesota 55455, and 8Department of Anatomy, Research Institute Neurosciences, Vrije Universiteit University Medical Center, 1007 MB Amsterdam, The Netherlands This mini-symposium aims to integrate recent insights from anatomy, behavior, and neurophysiology, highlighting the anatom- ical organization, behavioral significance, and information-processing mechanisms of corticostriatal interactions. In this sum- mary of topics, which is not meant to provide a comprehensive survey, we will first review the anatomy of corticostriatal circuits, comparing different ways by which “loops” of cortical–basal ganglia circuits communicate. Next, we will address the causal importance and systems-neurophysiological mechanisms of corticostriatal interactions for memory, emphasizing the communi- cation between hippocampus and ventral striatum during contextual conditioning. Furthermore, ensemble recording techniques have been applied to compare information processing in the dorsal and ventral striatum to predictions from reinforcement learning theory. -
Understanding the Building Blocks of Avian Complex Cognition : The
Understanding the building blocks of avian complex cognition: the executive caudal nidopallium and the neuronal energy budget by Kaya von Eugen A thesis submitted in partial fulfilment of the requirements for the degree of Philosophiae Doctoris (PhD) in Neuroscience From the International Graduate School of Neuroscience Ruhr University Bochum September 30th 2020 This research was conducted at the Department of Biopsychology, within the Faculty of Psychology at the Ruhr University under the supervision of Prof. Dr. Dr. h.c. Onur Güntürkün Printed with the permission of the International Graduate School of Neuroscience, Ruhr University Bochum Statement I certify herewith that the dissertation included here was completed and written independently by me and without outside assistance. References to the work and theories of others have been cited and acknowledged completely and correctly. The “Guidelines for Good Scientific Practice” according to § 9, Sec. 3 of the PhD regulations of the International Graduate School of Neuroscience were adhered to. This work has never been submitted in this, or a similar form, at this or any other domestic or foreign institution of higher learning as a dissertation. The abovementioned statement was made as a solemn declaration. I conscientiously believe and state it to be true and declare that it is of the same legal significance and value as if it were made under oath. Bochum, 30.09.2020 Kaya von Eugen PhD Commission Chair: PD Dr. Dirk Jancke 1st Internal Examiner: Prof. Dr. Dr. h.c. Onur Güntürkün 2nd Internal Examiner: Prof. Dr. Carsten Theiß External Examiner: Prof. Dr. Andrew Iwaniuk Non-Specialist: Prof. -
Emotional and Spatial Learning in Goldfish Is Dependent on Different
European Journal of Neuroscience, Vol. 21, pp. 2800–2806, 2005 ª Federation of European Neuroscience Societies Emotional and spatial learning in goldfish is dependent on different telencephalic pallial systems Manuel Portavella1,* and Juan P. Vargas2 1Departamento de Psicologı´a Experimental. Universidad de Sevilla. C ⁄ Camilo Jose´ Cela s ⁄ n, E-41018, Seville, Spain 2SISSA. Cognitive Neuroscience Sector. Via Beirut, 2 ⁄ 4, 34014 Trieste, Italy Keywords: amygdala, avoidance learning, brain evolution, hippocampus, memory systems Abstract In mammals, the amygdala and the hippocampus are involved in different aspects of learning. Whereas the amygdala complex is involved in emotional learning, the hippocampus plays a critical role in spatial and contextual learning. In fish, it has been suggested that the medial and lateral region of the telencephalic pallia might be the homologous neural structure to the mammalian amygdala and hippocampus, respectively. Although there is evidence of the implication of medial and lateral pallium in several learning processes, it remains unclear whether both pallial areas are involved distinctively in different learning processes. To address this issue, we examined the effect of selective ablation of the medial and lateral pallium on both two-way avoidance and reversal spatial learning in goldfish. The results showed that medial pallium lesions selectively impaired the two-way avoidance task. In contrast, lateral pallium ablations impaired the spatial task without affecting the avoidance performance. These results indicate that the medial and lateral pallia in fish are functionally different and necessary for emotional and spatial learning, respectively. Present data could support the hypothesis that a sketch of these regions of the limbic system, and their associated functions, were present in the common ancestor of fish and terrestrial vertebrates 400 million years ago. -
Cortical Layers: What Are They Good For? Neocortex
Cortical Layers: What are they good for? Neocortex L1 L2 L3 network input L4 computations L5 L6 Brodmann Map of Cortical Areas lateral medial 44 areas, numbered in order of samples taken from monkey brain Brodmann, 1908. Primary visual cortex lamination across species Balaram & Kaas 2014 Front Neuroanat Cortical lamination: not always a six-layered structure (archicortex) e.g. Piriform, entorhinal Larriva-Sahd 2010 Front Neuroanat Other layered structures in the brain Cerebellum Retina Complexity of connectivity in a cortical column • Paired-recordings and anatomical reconstructions generate statistics of cortical connectivity Lefort et al. 2009 Information flow in neocortical microcircuits Simplified version “computational Layer 2/3 layer” Layer 4 main output Layer 5 main input Layer 6 Thalamus e - excitatory, i - inhibitory Grillner et al TINS 2005 The canonical cortical circuit MAYBE …. DaCosta & Martin, 2010 Excitatory cell types across layers (rat S1 cortex) Canonical models do not capture the diversity of excitatory cell classes Oberlaender et al., Cereb Cortex. Oct 2012; 22(10): 2375–2391. Coding strategies of different cortical layers Sakata & Harris, Neuron 2010 Canonical models do not capture the diversity of firing rates and selectivities Why is the cortex layered? Do different layers have distinct functions? Is this the right question? Alternative view: • When thinking about layers, we should really be thinking about cell classes • A cells class may be defined by its input connectome and output projectome (and some other properties) • The job of different classes is to (i) make associations between different types of information available in each cortical column and/or (ii) route/gate different streams of information • Layers are convenient way of organising inputs and outputs of distinct cell classes Excitatory cell types across layers (rat S1 cortex) INTRATELENCEPHALIC (IT) | PYRAMIDAL TRACT (PT) | CORTICOTHALAMIC (CT) From Cereb Cortex. -
The Structural Model: a Theory Linking Connections, Plasticity, Pathology, Development and Evolution of the Cerebral Cortex
Brain Structure and Function https://doi.org/10.1007/s00429-019-01841-9 REVIEW The Structural Model: a theory linking connections, plasticity, pathology, development and evolution of the cerebral cortex Miguel Ángel García‑Cabezas1 · Basilis Zikopoulos2,3 · Helen Barbas1,3 Received: 11 October 2018 / Accepted: 29 January 2019 © Springer-Verlag GmbH Germany, part of Springer Nature 2019 Abstract The classical theory of cortical systematic variation has been independently described in reptiles, monotremes, marsupials and placental mammals, including primates, suggesting a common bauplan in the evolution of the cortex. The Structural Model is based on the systematic variation of the cortex and is a platform for advancing testable hypotheses about cortical organization and function across species, including humans. The Structural Model captures the overall laminar structure of areas by dividing the cortical architectonic continuum into discrete categories (cortical types), which can be used to test hypotheses about cortical organization. By type, the phylogenetically ancient limbic cortices—which form a ring at the base of the cerebral hemisphere—are agranular if they lack layer IV, or dysgranular if they have an incipient granular layer IV. Beyond the dysgranular areas, eulaminate type cortices have six layers. The number and laminar elaboration of eulaminate areas differ depending on species or cortical system within a species. The construct of cortical type retains the topology of the systematic variation of the cortex and forms the basis for a predictive Structural Model, which has successfully linked cortical variation to the laminar pattern and strength of cortical connections, the continuum of plasticity and stability of areas, the regularities in the distribution of classical and novel markers, and the preferential vulnerability of limbic areas to neurodegenerative and psychiatric diseases. -
Foxg1confines Cajal–Retzius Neuronogenesis and Hippocampal
The Journal of Neuroscience, April 27, 2005 • 25(17):4435–4441 • 4435 Development/Plasticity/Repair Foxg1 Confines Cajal–Retzius Neuronogenesis and Hippocampal Morphogenesis to the Dorsomedial Pallium Luca Muzio and Antonello Mallamaci Department of Biological and Technological Research, San Raffaele Scientific Institute, 20132 Milan, Italy It has been suggested that cerebral cortex arealization relies on positional values imparted to early cortical neuroblasts by transcription factor genes expressed within the pallial field in graded ways. Foxg1, encoding for one of these factors, previously was reported to be necessary for basal ganglia morphogenesis, proper tuning of cortical neuronal differentiation rates, and the switching of cortical neuro- blasts from early generation of primordial plexiform layer to late production of cortical plate. Being expressed along a rostral/lateral high- to-caudal/medial low gradient, Foxg1, moreover, could contribute to shaping the cortical areal profile as a repressor of caudomedial fates. Wetestedthispredictionbyavarietyofapproachesandfoundthatitwascorrect.WefoundthatoverproductionofCajal–Retziusneurons characterizing Foxg1Ϫ/Ϫ mutants does not arise specifically from blockage of laminar histogenetic progression of neocortical neuro- blasts, as reported previously, but rather reflects lateral-to-medial repatterning of their cortical primordium. Even if lacking a neocortical plate, Foxg1Ϫ/Ϫ embryos give rise to structures, which, for molecular properties and birthdating profile, are highly reminiscent of hippocampal plate and dentate blade. Remarkably, in the absence of Foxg1, additional inactivation of the medial fates promoter Emx2, although not suppressing cortical specification, conversely rescues overproduction of Reelin on neurons. Key words: Foxg1; Emx2; Wnt types; hippocampus; neocortex; Cajal–Retzius cells Introduction neuronogenesis (Xuan et al., 1995; Dou et al., 1999; Seoane et al., Areal specification of cortical neurons is an extremely complex 2004). -
The Ventral Striatum in Off-Line Processing: Ensemble Reactivation During Sleep and Modulation by Hippocampal Ripples
6446 • The Journal of Neuroscience, July 21, 2004 • 24(29):6446–6456 Behavioral/Systems/Cognitive The Ventral Striatum in Off-Line Processing: Ensemble Reactivation during Sleep and Modulation by Hippocampal Ripples C. M. A. Pennartz,1 E. Lee,1 J. Verheul,1 P. Lipa,2 C. A. Barnes,2 and B. L. McNaughton2 1Graduate School of Neurosciences Amsterdam, University of Amsterdam, Faculty of Science, Swammerdam Institute for Life Sciences, 1090 GB, Amsterdam, The Netherlands, and 2Arizona Research Laboratories Division of Neural Systems, Memory, and Aging, University of Arizona, Tucson, Arizona 85724 Previously it has been shown that the hippocampus and neocortex can spontaneously reactivate ensemble activity patterns during post-behavioral sleep and rest periods. Here we examined whether such reactivation also occurs in a subcortical structure, the ventral striatum, which receives a direct input from the hippocampal formation and has been implicated in guidance of consummatory and conditioned behaviors. During a reward-searching task on a T-maze, flanked by sleep and rest periods, parallel recordings were made from ventral striatal ensembles while EEG signals were derived from the hippocampus. Statistical measures indicated a significant amount of reactivation in the ventral striatum. In line with hippocampal data, reactivation was especially prominent during post- behavioralslow-wavesleep,butunlikethehippocampus,nodecayinpatternrecurrencewasvisibleintheventralstriatumacrossthefirst 40 min of post-behavioral rest. We next studied the relationship between ensemble firing patterns in ventral striatum and hippocampal ripples–sharp waves, which have been implicated in pattern replay. Firing rates were significantly modulated in close temporal associ- ation with hippocampal ripples in 25% of the units, showing a marked transient enhancement in the average response profile. -
Revised Nomenclature for Avian Telencephalon and Some Related Brainstem Nuclei
THE JOURNAL OF COMPARATIVE NEUROLOGY 473:377–414 (2004) Revised Nomenclature for Avian Telencephalon and Some Related Brainstem Nuclei ANTON REINER,1* DAVID J. PERKEL,2 LAURA L. BRUCE,3 ANN B. BUTLER,4 ANDRA´ S CSILLAG,5 WAYNE KUENZEL,6 LORETA MEDINA,7 GEORGE PAXINOS,8 TORU SHIMIZU,9 GEORG STRIEDTER,10 MARTIN WILD,11 GREGORY F. BALL,12 SARAH DURAND,13 ONUR GU¨ TU¨ RKU¨ N,14 DIANE W. LEE,15 CLAUDIO V. MELLO,16 ALICE POWERS,17 STEPHANIE A. WHITE,18 GERALD HOUGH,19 LUBICA KUBIKOVA,20 TOM V. SMULDERS,21 KAZUHIRO WADA,20 JENNIFER DUGAS-FORD,22 SCOTT HUSBAND,9 KEIKO YAMAMOTO,1 JING YU,20 CONNIE SIANG,20 AND ERICH D. JARVIS20* 1Department of Anatomy and Neurobiology, University of Tennessee Health Science Center, Memphis, Tennessee 38163 2Departments of Biology and Otolaryngology, University of Washington, Seattle, Washington 98195-6515 3Department of Biomedical Sciences, Creighton University School of Medicine, Omaha, Nebraska 68178 4Krasnow Institute and Department of Psychology, George Mason University, Fairfax, Virginia 22030-4444 5Department of Anatomy, Semmelweis University, Faculty of Medicine, H-1094, Budapest, Hungary 6Department of Poultry Science, Poultry Science Center, University of Arkansas, Fayetteville, Arkansas 72701 7Department of Human Anatomy, Faculty of Medicine, University of Murcia, Murcia E-30100, Spain 8Prince of Wales Medical Research Institute, Sydney, New South Wales 2031, Australia 9Department of Psychology, University of South Florida, Tampa, Florida 33620-8200 10Department of Neurobiology and Behavior, University -
REVIEW Hormones and the Hippocampus
205 REVIEW Hormones and the hippocampus R Lathe Centre for Genome Research and Centre for Neuroscience, University of Edinburgh, West Mains Road, Edinburgh EH9 3JQ, UK (Requests for offprints should be addressed to R Lathe, Centre for Genome Research, King’s Buildings, West Mains Road, Edinburgh EH9 3JQ, UK; Email: [email protected]) Abstract Hippocampal lesions produce memory deficits, but the Functionally, hippocampal enteroception may reflect feed- exact function of the hippocampus remains obscure. back control; evidence is reviewed that the hippocampus Evidence is presented that its role in memory may be modulates body physiology, including the activity of the ancillary to physiological regulation. Molecular studies hypothalamus–pituitary–adrenal axis, blood pressure, demonstrate that the hippocampus is a primary target for immunity, and reproductive function. It is suggested that ligands that reflect body physiology, including ion balance the hippocampus operates, in parallel with the amygdala, and blood pressure, immunity, pain, reproductive status, to modulate body physiology in response to cognitive satiety and stress. Hippocampal receptors are functional, stimuli. Hippocampal outputs are predominantly inhibi- probably accessible to their ligands, and mediate physio- tory on downstream neuroendocrine activity; increased logical and cognitive changes. This argues that an early synaptic efficacy in the hippocampus (e.g. long-term role of the hippocampus may have been in sensing soluble potentiation) could facilitate throughput inhibition. -
Broom Fish Brains Pain
Pre-publication copy Broom, D.M. 2016. Fish brains and behaviour indicate capacity for feeling pain. Animal Sentience, 2016.010 (5 pages). Fish brains, as well as fish behaviour, indicate capacity for awareness and feeling pain Donald M. Broom Centre for Anthrozoology and Animal Welfare Department of Veterinary Medicine University of Cambridge Madingley Road Cambridge CB3 0ES U.K. [email protected] http://www.neuroscience.cam.ac.uk/directory/profile.php?dmb16 Keywords pain sentience welfare fish feelings emotions brain behaviour Abstract Studies of behaviour are of major importance in understanding human pain and pain in other animals such as fish. Almost all of the characteristics of the mammalian pain system are also described for fish. Emotions, feelings and learning from these are controlled in the fish brain in areas anatomically different but functionally very similar to those in mammals. The evidence of pain and fear system function in fish is so similar to that in humans and other mammals that it is logical to conclude that fish feel fear and pain. Fish are sentient beings. Key (2015) is scornful about evidence from studies of fish behaviour indicating that fish are aware and feel pain but presents a thorough explanation of the pain system in the human brain and concludes that fish could not feel pain, or have any other feelings, as they do not have the brain structures that allow pain and other feelings in humans. Section 2 of his paper emphasises “the cortical origins of human pain” and states that “structure determines function”, eXplaining the functions of the five layers of the human cortex.