Theodore Holmes “Ted” Bullock (16 May 1915 – 20 December 2005) is one of the founding fathers of neuroethology

Neuroethology

Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system...

. During a career spanning nearly seven decades, this American academic was esteemed both as a pioneering and influential neuroscientist, examining the physiology and evolution of the nervous system across organizational levels, and as a champion of the comparative approach, studying species from nearly all major animal groups—coelenterates, annelids, arthropods, echinoderms, molluscs, and chordates.

Bullock discovered the pit organ in pit vipers and electroreceptors in weakly electric fish

Electric fish

An electric fish is a fish that can generate electric fields. It is said to be electrogenic; a fish that has the ability to detect electric fields is said to be electroreceptive. Most electrogenic fish are also electroreceptive. Electric fish species can be found both in the sea and in freshwater...

, as well as other electrosensory animals. His work on the jamming avoidance response

Jamming avoidance response

The jamming avoidance response or JAR is a behavior performed by some species of weakly electric fish. The JAR occurs when two electric fish with wave discharges meet – if their discharge frequencies are very similar, each fish will shift its discharge frequency to increase the difference between...

 in electric fish

Electric fish

An electric fish is a fish that can generate electric fields. It is said to be electrogenic; a fish that has the ability to detect electric fields is said to be electroreceptive. Most electrogenic fish are also electroreceptive. Electric fish species can be found both in the sea and in freshwater...

 (work later carried on by Walter Heiligenberg

Walter Heiligenberg

Walter Heiligenberg is best known for his contribution to neuroethology through his work on one of the best neurologically understood behavioral patterns in vertebrate, Eigenmannia...

) is an excellent example of how motor programs are integrated with incoming sensory information when generating a behavior pattern in response to a stimulus.

Bullock appealed to the scientific community to look beyond established paradigms in neuroscience, as well as to consider the ecology of an animal when endeavoring to understand its nervous system. As he once wrote, “Neuroscience is part of biology, more specifically of zoology, and it suffers tunnel vision unless continuous with ethology

Ethology

Ethology is the scientific study of animal behavior, and a sub-topic of zoology....

, ecology

Ecology

Ecology is the scientific study of the relations that living organisms have with respect to each other and their natural environment. Variables of interest to ecologists include the composition, distribution, amount , number, and changing states of organisms within and among ecosystems...

, and evolution

Evolution

Evolution is any change across successive generations in the heritable characteristics of biological populations. Evolutionary processes give rise to diversity at every level of biological organisation, including species, individual organisms and molecules such as DNA and proteins.Life on Earth...

”.

In his quest to go beyond a descriptive account of the nervous system, Bullock studied many different and
unrelated, species. He believed that this “comparative approach” would reveal both general principles of the nervous system, and offer insights into which nervous system properties (anatomical, physiological, and chemical) were relevant to observed differences in species-specific traits, as well as which specific traits were relevant to observed differences in nervous systems. His resulting discoveries helped explain various properties of nervous systems. In one influential review he wrote, “Comparative neuroscience is likely to reach insights so novel as to constitute revolutions in understanding the structure, functions, ontogeny, and evolution of nervous systems. […] Without due consideration of the neural and behavioral correlates of differences between higher taxa and between closely related families, species, sexes , and stages, we cannot expect to understand nervous systems or ourselves”.

One colleague described Bullock as an “adventurous scientific explorer, continually seeking undiscovered phenomena and new unifying principles”. Until the very end of his life, at the age of 90, Bullock remained an active and influential presence in the fields of neuroscience

Neuroscience

Neuroscience is the scientific study of the nervous system. Traditionally, neuroscience has been seen as a branch of biology. However, it is currently an interdisciplinary science that collaborates with other fields such as chemistry, computer science, engineering, linguistics, mathematics,...

 and neuroethology

Neuroethology

Neuroethology is the evolutionary and comparative approach to the study of animal behavior and its underlying mechanistic control by the nervous system...

.

Biography

The second of four children, Bullock was born May 16, 1915 in Nanking, China. His parents, Amasa and Ruth Bullock (née Beckwith), were Presbyterian missionaries and had arrived in China in 1909. In 1928, when Bullock was thirteen, the family returned to the United States, and settled in Southern California

Southern California

Southern California is a megaregion, or megapolitan area, in the southern area of the U.S. state of California. Large urban areas include Greater Los Angeles and Greater San Diego. The urban area stretches along the coast from Ventura through the Southland and Inland Empire to San Diego...

. Bullock’s life as a neuroscientist began with histological studies of brain degeneration that he performed while still in high school. During this time he also studied marine biology and other courses at the Pomona College

Pomona College

Pomona College is a private, residential, liberal arts college in Claremont, California. Founded in 1887 in Pomona, California by a group of Congregationalists, the college moved to Claremont in 1889 to the site of a hotel, retaining its name. The school enrolls 1,548 students.The founding member...

 Marine Biological Laboratory. He received an Associate of Arts degree from Pasadena Junior College in 1934, and a BA from University of California, Berkeley

University of California, Berkeley

The University of California, Berkeley , is a teaching and research university established in 1868 and located in Berkeley, California, USA...

 in 1936, where he studied zoology. In 1937 Bullock married Martha Runquist, to whom he would be with until the end of his life, 68 years later. They would have two children, Christine and Steve.

Bullock’s doctorate work was performed at UC Berkeley under the supervision of S.F. Light, and focused on the organization of the nervous system (both anatomy and physiology) in acorn worm

Acorn worm

The Acorn worms or Enteropneusta are a hemichordate class of invertebrates, closely related to the echinoderms. There are about 90 species of acorn worm in the world, the main species for research being Saccoglossus kowaleskii....

s, generally considered a sister group to the chordates. This marked the beginning of his studies on simple nervous systems, which he used to explore the neural mechanisms that work together to produce an output in response to a stimulus, both at the physiological and behavioral level. During this time, the importance of comparative studies also became apparent to him. He believed that to fully understand how the brain and nervous system work, one must search for commonalities, and also for differences in nervous systems across different taxonomic levels.
After earning his PhD in 1940, he accepted a postdoctoral fellowship, and later a teaching position at Yale

YALE

RapidMiner, formerly YALE , is an environment for machine learning, data mining, text mining, predictive analytics, and business analytics. It is used for research, education, training, rapid prototyping, application development, and industrial applications...

. During his four years at Yale

YALE

RapidMiner, formerly YALE , is an environment for machine learning, data mining, text mining, predictive analytics, and business analytics. It is used for research, education, training, rapid prototyping, application development, and industrial applications...

, Bullock worked at the Marine Biological Laboratory

Marine Biological Laboratory

The Marine Biological Laboratory is an international center for research and education in biology, biomedicine and ecology. Founded in 1888, the MBL is the oldest independent marine laboratory in the Americas, taking advantage of a coastal setting in the Cape Cod village of Woods Hole, Massachusetts...

 (MBL) at Woods Hole during the summers. Here he taught invertebrate zoology and their famous physiology course, and he studied nerve nets in coelenterates and the structure and physiology of giant nerve fibers in annelids. His studies on nerve nets lead him to be one of the first experimentalists to understand the value and importance of computational techniques for modeling and data analysis.

In 1944 Bullock accepted a faculty position at the University of Missouri

University of Missouri

The University of Missouri System is a state university system providing centralized administration for four universities, a health care system, an extension program, five research and technology parks, and a publishing press. More than 64,000 students are currently enrolled at its four campuses...

, where he taught medical students anatomy and physiology. Two years later he joined the faculty at University of California, Los Angeles

University of California, Los Angeles

The University of California, Los Angeles is a public research university located in the Westwood neighborhood of Los Angeles, California, USA. It was founded in 1919 as the "Southern Branch" of the University of California and is the second oldest of the ten campuses...

 (UCLA), where he remained for the next twenty years. During this time, he helped pioneer the field of comparative and integrative neurobiology. In one series of famous experiments on the cardiac ganglion in lobsters, Bullock demonstrated that neurons can communicate not just via action potential

Action potential

In physiology, an action potential is a short-lasting event in which the electrical membrane potential of a cell rapidly rises and falls, following a consistent trajectory. Action potentials occur in several types of animal cells, called excitable cells, which include neurons, muscle cells, and...

 and chemical synapse

Chemical synapse

Chemical synapses are specialized junctions through which neurons signal to each other and to non-neuronal cells such as those in muscles or glands. Chemical synapses allow neurons to form circuits within the central nervous system. They are crucial to the biological computations that underlie...

, but through non-synaptic interactions without such impulses. Today we know that this type of electrical interaction is mediated by gap junctions. This idea, that electrical synapses couple groups of cells into functional units, lead to Bullock’s lifelong interest in field potentials

Local field potential

A local field potential is a particular class of electrophysiological signals, which is dominated by the electrical current flowing from all nearby dendritic synaptic activity within a volume of tissue. A voltage is produced by the summed synaptic current flowing across the resistance of the local...

, which are generated by the summated electrical activity of millions of brain cells. Bullock was a respected teacher who taught many courses while at UCLA, such as zoology and advanced invertebrate biology. He spent the summers of 1955-1957 at Woods Hole as the director of the Invertebrate Zoology course.

In 1966 Bullock left UCLA to join the University of California, San Diego

University of California, San Diego

The University of California, San Diego, commonly known as UCSD or UC San Diego, is a public research university located in the La Jolla neighborhood of San Diego, California, United States...

 (UCSD) School of Medicine’s new Department of Neurosciences. He also served as the chairman of the Neurobiology Unit of Scripps Institution of Oceanography

Scripps Institution of Oceanography

Scripps Institution of Oceanography in La Jolla, California, is one of the oldest and largest centers for ocean and earth science research, graduate training, and public service in the world...

 in La Jolla, CA. One reason for his move to UCSD was that he hoped to bridge the gap between Marine Biology and medicine.

Bullock published a vast array of papers. Other than the species previously mentioned, he also studied the nervous systems of coral

Coral

Corals are marine animals in class Anthozoa of phylum Cnidaria typically living in compact colonies of many identical individual "polyps". The group includes the important reef builders that inhabit tropical oceans and secrete calcium carbonate to form a hard skeleton.A coral "head" is a colony of...

s, sea urchin

Sea urchin

Sea urchins or urchins are small, spiny, globular animals which, with their close kin, such as sand dollars, constitute the class Echinoidea of the echinoderm phylum. They inhabit all oceans. Their shell, or "test", is round and spiny, typically from across. Common colors include black and dull...

s, spirunculids, Limulus, Aplysia

Aplysia

Aplysia is a genus of medium-sized to extremely large sea slugs, specifically sea hares, which are one clade of large sea slugs, marine gastropod mollusks. The general description of sea hares can be found in the article on the superfamily Aplysioidea....

, starfish, rattlesnake

Rattlesnake

Rattlesnakes are a group of venomous snakes of the genera Crotalus and Sistrurus of the subfamily Crotalinae . There are 32 known species of rattlesnake, with between 65-70 subspecies, all native to the Americas, ranging from southern Alberta and southern British Columbia in Canada to Central...

s, rays

Batoidea

Batoidea is a superorder of cartilaginous fish commonly known as rays and skates, containing more than 500 described species in thirteen families...

, shark

Shark

Sharks are a type of fish with a full cartilaginous skeleton and a highly streamlined body. The earliest known sharks date from more than 420 million years ago....

s, porpoise

Porpoise

Porpoises are small cetaceans of the family Phocoenidae; they are related to whales and dolphins. They are distinct from dolphins, although the word "porpoise" has been used to refer to any small dolphin, especially by sailors and fishermen...

s, sea lion

Sea Lion

Sea lions are pinnipeds characterized by external ear-flaps, long fore-flippers, the ability to walk on all fours, and short thick hair. Together with the fur seal, they comprise the family Otariidae, or eared seals. There are six extant and one extinct species in five genera...

s, cuttlefish

Cuttlefish

Cuttlefish are marine animals of the order Sepiida. They belong to the class Cephalopoda . Despite their name, cuttlefish are not fish but molluscs....

, catfish

Catfish

Catfishes are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the heaviest and longest, the Mekong giant catfish from Southeast Asia and the second longest, the wels catfish of Eurasia, to detritivores...

, sloth

Sloth

Sloths are the six species of medium-sized mammals belonging to the families Megalonychidae and Bradypodidae , part of the order Pilosa and therefore related to armadillos and anteaters, which sport a similar set of specialized claws.They are arboreal residents of the jungles of Central and South...

s, manatee

Manatee

Manatees are large, fully aquatic, mostly herbivorous marine mammals sometimes known as sea cows...

s, salamander

Salamander

Salamander is a common name of approximately 500 species of amphibians. They are typically characterized by a superficially lizard-like appearance, with their slender bodies, short noses, and long tails. All known fossils and extinct species fall under the order Caudata, while sometimes the extant...

s, frog

Frog

Frogs are amphibians in the order Anura , formerly referred to as Salientia . Most frogs are characterized by a short body, webbed digits , protruding eyes and the absence of a tail...

s, turtle

Turtle

Turtles are reptiles of the order Testudines , characterised by a special bony or cartilaginous shell developed from their ribs that acts as a shield...

s, hagfish

Hagfish

Hagfish, the clade Myxini , are eel-shaped slime-producing marine animals . They are the only living animals that have a skull but not a vertebral column. Along with lampreys, hagfish are jawless and are living fossils whose next nearest relatives include all vertebrates...

, crayfish

Crayfish

Crayfish, crawfish, or crawdads – members of the superfamilies Astacoidea and Parastacoidea – are freshwater crustaceans resembling small lobsters, to which they are related...

, tuna

Tuna

Tuna is a salt water fish from the family Scombridae, mostly in the genus Thunnus. Tuna are fast swimmers, and some species are capable of speeds of . Unlike most fish, which have white flesh, the muscle tissue of tuna ranges from pink to dark red. The red coloration derives from myoglobin, an...

, ratfish, bat

Bat

Bats are mammals of the order Chiroptera "hand" and pteron "wing") whose forelimbs form webbed wings, making them the only mammals naturally capable of true and sustained flight. By contrast, other mammals said to fly, such as flying squirrels, gliding possums, and colugos, glide rather than fly,...

s, crab

Crab

True crabs are decapod crustaceans of the infraorder Brachyura, which typically have a very short projecting "tail" , or where the reduced abdomen is entirely hidden under the thorax...

s, octopodes, snake

Snake

Snakes are elongate, legless, carnivorous reptiles of the suborder Serpentes that can be distinguished from legless lizards by their lack of eyelids and external ears. Like all squamates, snakes are ectothermic, amniote vertebrates covered in overlapping scales...

s, rat

Rat

Rats are various medium-sized, long-tailed rodents of the superfamily Muroidea. "True rats" are members of the genus Rattus, the most important of which to humans are the black rat, Rattus rattus, and the brown rat, Rattus norvegicus...

s and human

Human

Humans are the only living species in the Homo genus...

s. In 1965 together with Adrian Horridge, Bullock published the seminal two-volume “bible of invertebrate neurobiology”: Structure and Function in the Nervous System of Invertebrates.

Bullock was known as an inspired teacher and mentor. More than 100 scientists passed through his laboratory as postdoctoral fellows and research associates. From 1949 to 1999, Bullock was the primary adviser for 36 graduating PhD students (17 at Scripps), and in 1982 he retired as a Professor Emeritus. However, retirement could not stop him from remaining at the forefront of comparative neuroscience. At the age of 88 Bullock re-established a modeling study on nerve-nets, and built a model that accurately predicted the input-output relationships for a range of different stimuli. Bullock maintained an active research laboratory and continued studying the anatomy and physiology of the nervous system up until his death, 20 December 2005.

Notable awards and positions

1950–51, Fulbright scholarship, The Zoological Station, Naples, Italy

1955–56, President, The American Association of University Professors

American Association of University Professors

The American Association of University Professors is an organization of professors and other academics in the United States. AAUP membership is about 47,000, with over 500 local campus chapters and 39 state organizations...

1961, Elected as a member, The America Academy of Arts and Science

1963, Admitted into the National Academy of Sciences

United States National Academy of Sciences

The National Academy of Sciences is a corporation in the United States whose members serve pro bono as "advisers to the nation on science, engineering, and medicine." As a national academy, new members of the organization are elected annually by current members, based on their distinguished and...

 (NAS), served as chair of the NAS Zoology Section, and when it was later dissolved he became chair of the new Section of Neurobiology

1965, President, The American Society of Zoologists (now the Society for Integrative and Comparative Biology

Society for Integrative and Comparative Biology

The Society for Integrative and Comparative Biology is organized to integrate the many fields of specialization which occur in the broad field of biology. The society is dedicated to promoting the pursuit and dissemination of important information relating to biology...

 (SICB))

1968, Karl Spencer Lashley Award, The American Philosophical Society

1970, Elected as a member, The American Philosophical Society

1973, Queen’s Fellow in Marine Biology, Australia

1973-4, Third president of The Society for Neuroscience

1984, First president of the International Society for Neuroethology

1984, Ralph W. Gerard Prize, The Society for Neuroscience

1988, Honorary Doctorate, University of Frankfurt

2000, Honorary Doctorate, University of Loyola Chicago

More information

• Bullock’s UCSD profile page, with a list of his publications: http://myprofile.cos.com/bullockt82s
• Bullock's autobiography: http://cogprints.org/130/0/Autobiog.htm