FISIOLOGIA HUMANA

Dr. NÉSTOR RODRIGUEZ ALAYO

[email protected]
 BIOLOGIA
MOLECULAR
BIOLOGIA
BIOQUIMICA CELULAR

FISIOLOGIA FARMACOLOGIA
BIOMATEMATICA

ANATOMIA
INMUNOLOGIA
GENETICA
 FISIOLOGIA
• Estudio de la vida y como funciona los genes, células,
tejidos y organismos.
• «Integrando ciencias de la vida, desde lo molecular al
organismo: su adaptación»
• «La investigación animal salva vidas humanas»
• Genómica – Proteómica - Fisionómica
• Proyecto Cerebro Humano (2013-2016) descifrar función
topográfica e integrativa cerebral: Neurociencias.
• Nueva hipótesis de la fiebre.
Beverly Bishop
October 19, 1922 - September 20, 2008
The broad objective of my research over the
years, has been to identify and analyze how the
nervous system controls muscle activity in the
intact behaving animal including human subjects.
Much of my experimental work has been
concerned with the neural regulation of the
respiratory muscles. My major contribution in this
area was defining the neural pathways of the
reflex circuits which control the expiratory activity
of the abdominal muscles, the major expiratory
muscles in vertebrates. Work in this area is still
going on.
Clark Blatteis
June 25, 1932, Berlin, Germany
Clark Blatteis, professor of physiology at the University
of Tennessee Health Science Center, is interviewed in
this segment of “Living History of Physiology” by APS
Executive Director Dr. Martin Frank. Dr. Blatteis has an
outstanding career of more than 50 years focusing on
the pathophysiology of fever and how the body’s
physiological processes respond to noxious effects of
infectious agents.
Mordecai P. Blaustein

Mordecai Blaustein, a professor of physiology and

medicine at the University of Maryland School of Medicine, grew
up in Brooklyn, NY.  He received his first chemistry set at an early
age and, as a result, became enamored with the idea of
becoming a scientist (before he really knew what a scientist
was).  As an undergraduate at Cornell University (B.A. with
honors in zoology), he was introduced to the excitement of
physiology and research by Howard Schneiderman, and to
neurophysiology by William Van der Kloot.  During his medical
student days at Washington University in St. Louis, Blaustein
received NIH training funds to support a research project on
the sodium pump (Na, K, ATPase) with Daniel Tosteson
Felix Bronner
November 7, 1921, Vienna, Austria
Felix Bronner, retired professor of physiology in
biophysics and current editor of Topics of Bone Biology,
is interviewed in this segment of “Living History of
Physiology” by Carolyn Pennington of the Office of
Communications at the University of Connecticut Health
Center. Dr. Bronner has had a career of more than 50
years working on the physiology of calcium and bone
metabolism, as well as nutrition. He officially retired in
1989, but he is still active in research, writing and
editing, and organizing talks and seminar at the
University of Connecticut.
Maurice Burg , MD
Maurice Burg, principle investigator at the Lab of Kidney
and Electrolyte Metabolism, is interviewed in 2005 in this
segment of “Living History of Physiology” by Mark
Knepper, Chief of the Lab of Kidney and Electrolyte
Metabolism. Dr. Burg has had a career that spans nearly
50 years, as he has been a leader in the field of renal
physiology. He is known for his development of critical
technology that has led to new pathways of discovering
physiological principles that help us understand renal
function and osmotic regulation. He is perhaps best
known for his invention of the isolated perfused tubule
technique. 
Elsworth Buskirk
August 11, 1929 - March 28, 2010
Elsworth R. Buskirk received a B.A. in biology and
physical education from St. Olaf College, a master's
degree in physical education from the University of
Minnesota, and, in 1954, a Ph.D. in physiology, also
from the University of Minnesota. After completing
his Ph.D., Buskirk worked as chief of the
Environmental Physiology Section at the
Quartermaster Research and Development Center in
Natick, Massachusetts. From 1957 to 1963, he held
the position of research physiologist at the National
Institutes of Health, and, in 1963, he became a
faculty member in the Department of Physiology at
the Pennsylvania State University.
Oscar A. Candia
In a June 2005 letter published in The
Physiologist (48(3)) under “Senior
Physiologists’ News,” Oscar A. Candia wrote:
“I am continuing with my scientific activities. I
am a Professor of Ophthalmology and
Professor of Physiology here at the Mount
Sinai School of Medicine in New York where I
have been since 1968. I am also Vice-chairman
and Director of Research in the Department of
Ophthalmology.
Shu Chien
Shu Chien was installed as the 63th President of the
American Physiological Society at the close of the Society's
spring meeting in Washington, DC.
Chien, a professor of bioengineering and medicine at the
University of California, San Diego, succeeds Vernon S.
Bishop.
A 43-year member of APS, Chien has served on the
editorial boards of the American Journal of Physiology, the
Journal of Applied Physiology, and the American Journal of
Physiology: Heart and Circulatory Physiology. He also has
been an active contributor to the Society's educational
programs, including the slide-tape program on peripheral
circulation and the Handbook of Physiology: Circulation. In
1985, he was elected to the APS Council by the
membership and last ear was elected President-elect.
Jerry Clayton Collins received the Bachelor
of Engineering degree from Vanderbilt
University in 1962, the Master of Science
degree from Purdue University in 1965, and
the Ph.D. from Duke University in 1970, all in
electrical engineering. He and his wife Sandra
are the parents of three children: Leslie and
Reid, physicians, and Erin, an artist and
teacher, and grandparents of Isaac, Rachel, and
Silas.
Professor Collins worked at the U.S. Naval
Ordnance Laboratory in Silver Spring Maryland
and had teaching assignments in aviation
electronics at Purdue and in electrical
engineering at Duke before moving to the
University of Kentucky in 1968, where he
worked in the Department of Electrical
Engineering and the Division of Cardiothoracic
Surgery.
Helen Cooke
Helen Cooke was born in Greenfield,
Massachusetts in 1943.  She received a B.S.
degree from the University Massachusetts in
1965 and an M.S. from U.C.L.A. in 1967.  Shortly
thereafter, she moved to Australia and obtained a
Ph.D. in 1971 from the University of Sydney on
the topic of electrolyte and amino acid transport
in the kidney.  Dr. Cooke returned to the United
States and accepted a part-time appointment in
1971 as Instructor in the Physiology Department
at the University of Iowa Medical School. 
Robert Kellogg Crane
(December 20, 1919 - October 31, 2010)
As a rock-ribbed, dyed in the wool biochemist I was
surprised, pleased and felt greatly honored to be
included in the Living History program together with
such a distinguished group of physiologists.
It came about because:
(1) On April 1, 1966, I journeyed back to my
homeland, New Jersey, to become the founding
Chairman of the Department of Physiology and
Biophysics) at Rutgers University Medical School,
now the Robert Wood Johnson Medical School, a
component of the University of Medicine and
Dentistry of New Jersey. On assuming the post, I felt
comfortable with my degree from Harvard since it
was in Medical Science, not specifically in
Biochemistry, the same as it would have been had
my major been Physiology.  Nonetheless, I quickly
sought some more specific credentials and asked,
first, for membership in The Society of General
Physiologists and then in The American Physiological
Society. 
John Faulkner
John Faulkner's research focuses on the
contractile properties of whole skeletal
muscles, motor units, and single skeletal
muscle fibers from mice and rats and single
fibers from muscle biopsies of humans. In
particular, he and his associates are interested
in the changes that occur in the contractile
properties of skeletal muscle fibers during and
following injury, fatigue and regeneration.
Skeletal muscle fibers are injured by a variety
of diseases; local anesthetics; direct injury,
such as crushing and cutting, or puncturing;
extremes of temperature; ischemia; and
development of high forces during
contractions.
G. Edgar Folk, Jr.
G. Edgar Folk, Professor Emeritus of Molecular
Physiology at the University of Iowa, is
interviewed in July 2007 in this segment of
“Living History of Physiology” by Charles
Wunder, also a Professor Emeritus at the
University of Iowa. Dr. Folk’s career has
focused largely on hibernation and circadian
rhythm/biological clocks. He was one of the
founders of the fact that indeed, bears do
hibernate during the cold months.
MartinFrank

I was born in Chicago, Illinois and grew up with

the natural curiosity of youth, exploring the
world around me. That led me to science and
the associated science projects and science
fairs that one pursues while growing up. For
me, that interest translated into an interest in
pursuing a degree in science at the University
of Illinois, Urbana-Champaign. The question
was how could my parents afford to send me
to college, the first in my family to attend.
Fortunately, one of my summer jobs was to
work as a caddie at Bryn Mawr Country Club in
Lincolnwood, IL. It was while caddying that I
discovered that the Western Golf Association
had a scholarship program for caddies in
financial need. I attended college on an Evans
Scholarship that provided tuition and housing
in the Evans Scholars House on campus.
Gerhard Giebisch, M.D., is Sterling Professor
Emeritus of Cellular and Molecular Physiology at the
Yale University School of Medicine.  He was born and
raised in Vienna, Austria, where he attended
elementary and high school and received the M.D.
degree from the University of Vienna in 1951.  Dr.
Giebisch’s original career goal was to be an internist,
and he was advised by one of his professors, Erwin
Deutsch, to first get basic science research training.  Dr.
Deutsch referred this eager medical student to Franz
von Brücke, the Chairman of Pharmacology, who
greatly stimulated the young man’s interest in
physiology.  While still a student, Dr. Giebisch was first
introduced to renal physiology by reading a copy of
Homer Smith’s Porter Lectures that he had been given. 
He also read a book by Otto Spühler on modern
methods to study renal function, which led him to
contact Dr. Spühler and arrange for a three-month
period in his laboratory in Zurich. 
Malcolm S. Gordon 
Malcolm Gordon was born in Brooklyn, New
York City, November 13, 1933.  Growing up he
and his family lived in several places in
Brooklyn, northern New Jersey and Los
Angeles, CA.  He earned his undergraduate
degree in zoology from Cornell University in
1954.  He earned his PhD in zoology from Yale
University in 1958.  He did most of his doctoral
research at the Woods Hole Oceanographic
Institution, Woods Hole, Massachusetts.  He
spent a postdoctoral year (1957-8) in the
Department of Zoology, University of
Cambridge, U.K., then joined the faculty in the
then Department of Zoology (now Department
of Ecology and Evolutionary Biology) at the
University of California, Los Angeles (UCLA).
John E. Greenleaf
John E. Greenleaf, former research
physiologist at the NASA Ames Research
Center near Stanford University, is interviewed
in September 2007 in this segment of “Living
History of Physiology” by Donald Watenpaugh,
Ph.D. Dr. Greenleaf established and directed
the Laboratory for Human Environmental
Physiology in the Life Sciences Division at the
research center. Over his 40-year career, he has
had 181 published research articles, 127
abstracts, 23 book chapters, 75 other
publications and numerous awards.
William Hansel

Education:
M.S., Cornell University, New York, NY, 1947,
Animal Physiology
Ph.D., Cornell University, New York, NY, 1949,
Animal Physiology
Research Interests:
Dr. Hansel's major research interests are in
cancer, reproductive biology and nutrition. He
currently leads a group of scientists at the
Pennington Center and on the LSU Campus
that is developing a new method for treating
prostate, breast, testes, and ovarian cancers
based on targeting lytic peptides to
gonadotropin receptors on the cancer cell
membranes. Selected Publications:
Leuschner C, Hansel W. Membrane disrupting
lytic peptides for cancer treatments. Curr
Pharm Des. 2004;10(19):2299-310
Paul C. Johnson

Paul C. Johnson has been a member of the

American Physiological Society since 1959.  At
the time of the Living History interview,
Johnson was Professor Emeritus, University of
Arizona, Department of Physiology and Adjunct
Professor, University of California, San Diego,
Department of Bioengineering.  Paul Johnson
went to high school in Ironwood, Michigan and
pursued his B.S (physics) and Ph.D. (physiology)
degrees at the University of Michigan, Ann
Arbor.  Upon completion of his degree in 1955,
he became an instructor in the Department of
Physiology at the University of Michigan.  From
1956-1958, he was an instructor at Western
Reserve University in the Department of
Physiology
Gabor Kaley
November 16, 1926, Hungary
"I don't think about retiring. Intellectually, I feel I'm at
the top of my game and I don't believe I've even slowed
down much."
To call Gabe Kaley an elder statesman would probably
insult him, but only the elder part. Vigorous and
energetic, he treats his age as irrelevant: “In the last five
years, I have been more productive and have published
more papers than in any similar period of my life.” Not
bad for the dedicated, dapper professor who is chairman
of the Department of Physiology. As for retirement, if it
happens Dr. Kaley will have left his mark profoundly on
the study of physiology and on New York Medical College,
both of which claim his intensive devotion. And as the
longest sitting chair of physiology in the nation, he’s
succeeded by virtue of his reputation, scientific
accomplishments and the steadfast conviction that
physiology, as the science of living things, is the very basis
of medicine.
Arnold M. Katz
Dr. Katz was born in Chicago on July 30, 1932. His mother,
a graduate of the Cleveland Conservatory of Music, was a
piano teacher. Louis N. Katz, his father, was an
internationally renowned cardiologist and basic science
investigator who received a Lasker award, and was
President of the American Physiological Society and
American Heart Association.
Dr. Arnold Katz received his early education at the
University of Chicago Laboratory Schools, and graduated
from the University of Chicago College in 1952 earning a
B.A. with Honors in the Natural Sciences. While a college
student he studied yeast metabolism during the summer
of 1951 and spent the summer of 1952 at the Marine
Biological Laboratory in Woods Hole, MA where he
participated in a study of the physicochemical properties
of muscle. While a medical student at Harvard he spent
the summers of 1953 and 1954 with his father in Chicago
studying coronary flow, left ventricular volume curves and
the energetics of the heart.
Franklyn G. Knox

Franklyn G. Knox was born in Rochester, New York,

and completed all his professional education at
the now State University of New York at Buffalo.
He received the B.S. degree in 1959 and the M.D.
and Ph.D. degrees in 1965. The last of these
represented training in the Department of
Physiology and led to a position as staff associate
at the National Heart Institute (1965-68). From
there he moved to the University of Missouri,
where he was promoted to an associate
professorship in the Department of Physiology in
1970
Joan E. Lafuze

Joan Esterline Lafuze has been a member of

the American Physiological Society since she
was a graduate student in the 1970’s.  At the
time of the Living History Interview, Lafuze was
a Professor of Biology at Indiana University
East, Richmond, Indiana and Research and
Teaching Associate at Indiana University School
of Medicine in Indianapolis.  Joan Esterline
Lafuze graduated from Shortridge High School
in Indianapolis in 1955, Indiana University
Bloomington in 1959, St. Vincent Hospital
School of Medical Technology and was
registered with the American Society of
Medical Technologists in 1963
Samuel Leonard
(November 26, 1905 - November 12, 2007)
Dr. Samuel Leonard, Professor Emeritus of reproductive
physiology at Cornell University, is recorded at a
seminar at Cornell in November 1997. Dr. Leonard was
a pioneer in the fields of endocrinology and
reproductive physiology. His research focused on
hormones of the pituitary gland and thyroid gland, the
regulation of the mammary gland, and anywhere else
hormones are found. He taught introductory to zoology
and advanced endocrinology. At the age of 17, he
claims he was already excited by the discovery of
insulin, and by the age of 26, he had already published
seven papers and was a major player in endocrinology.
By 1931, Dr. Leonard had proven that estrogen could be
used as a contraceptive agent, and that the ovaries and
testes were regulated by two pituitary hormones –
follicle-stimulating hormone (FSH) and luteinizing
hormone (LH).
Richard Malvin
Dr. Richard Malvin, Professor Emeritus at the
University of  Michigan, is interviewed in this
October 2010 in this segment of "Living History
of Physiology" by APS Executive Director Dr.
Martin Frank. Dr. Malvin has been a member of
the American Physiological Society since 1960,
and a faculty member at the University of
Michigan since 1956. He has had a very
illustrious research career, and he is probably
best known for his development of the stop-flow
technique to measure renal function. Dr. Malvin
has published more than 150 papers.
Charles Rawlinson Park
Charles Rawlinson Park, better known as Rollo by his
many friends and associates, was born in Baltimore,
Maryland on March 2, 1916. His father, Edwards A.
Park, was on the staff of Johns Hopkins Medical School
and later became Chairman of the Department of
Pediatrics there. His mother introduced and supervised
nursery schools in Baltimore and subsequently
throughout Baltimore and Delaware.  Rollo graduated
A.B. from Harvard College in 1937 and M.D. in 1941
from Johns Hopkins, where he was elected to Alpha
Omega Alpha and Phi Beta Kappa.  He interned at Johns
Hopkins Hospital and then was appointed Resident and
Chief Resident in - Medicine at Peter Bent Brigham
Hospital in Boston.  During his student days at Johns
Hopkins, he made an auspicious entry in the world of
investigative medicine by reporting, in collaboration
with W. Barry Wood, Jr., that p-aminobenzoic acid was
an essential factor for bacterial growth.
Robert W. Phillips

Robert W. Phillips (Bob) was born in Peoria,

Illinois, in 1929. He was an unmotivated
student and after an unremarkable education
through high school in Peoria he spent several
lackadaisical years at Bradley and Kansas State
Universities. In June of 1950 he opted for a
change in venue and moved to Montana to
work on a cattle ranch. He arrived in June of
1950. The Korean conflict began that August.
He was near the top of the potential draftee
list and was soon inducted into the infantry.
Ananda S. Prasad

Ananda S. Prasad was born in Buxar, a small

town in the state of Bihar, India.  After
graduation from high school, he joined Patna
Science College, Patna University where he was
top student in chemistry and received B.Sc
degree with honors in mathematics.  Prasad
entered the Patna Medial College in Bihar,
graduating there in 1951 with high distinction
in physiology. 
Peter B. Raven
Peter B. Raven was born and raised in
London, England and immigrated to the
United States in 1965 to study at the
University of Oregon. He graduated with a
Ph.D. in the Scientific Basis of Physical
Education in 1969 and joined Steven M.
Horvath at the Institute of Environmental
Stress at the University of California at
Santa Barbara for a post-doctoral
fellowship. He joined the American
Physiological Society in 1971.
Loring Rowell
Loring (Larry) Rowell was born in Lynn,
Massachusetts on January 27, 1930.  He
graduated from high school in 1948 and
entered the University of Massachusetts,
Amherst, Massachusetts  in 1948 as a major in
Science and Biology.  Other interests were
skiing, football and track and field.  He
transferred to Springfield College in
Massachusetts in 1950 to prepare for a career
in teaching and coaching.   There, incidentally,
he met Charles Tipton, also the subject of a
video in this series, who became a life-long
friend and colleague.
Rafael Rubio
Rafael Rubio, professor at Universidad Autonoma de
San Luis Potosi in Mexico and emeritus professor at the
University of Virginia, is interviewed in January 2010 in
this segment of “Living History of Physiology” by Dr.
Maureen Knabb. Dr. Rubio has been a professor at
UASLP for 13 years. His undergraduate work was done
in Mexico City and only influenced by Dr. Arturo
Rosenblueth and Walter B. Cannon, for whom he was a
lab technician working on cardiovascular research. Dr.
Rubio has published almost 200 articles and book
chapters in his life, 25 of those coming before he even
obtained a bachelor’s degree. He was a full professor of
physiology at the University of Virginia until 1996 when
he returned to Mexico, where he is a professor of
physiology at UASLP. His research focuses on the
mechanism of coupling cardiac function and coronary
blood flow.
Born in Mexico during a time of civil war, his family
eventually
Bodil Schmidt-Nielsen
Bodil Schmidt-Nielsen brought to the presidency
of APS traditions different from those of her
recent predecessors. Of the eleven presidents
beginning with Pappenheimer (1964-65), seven
of them received a major part of their education
or training in laboratories at Harvard University.
Schmidt-Nielsen's background, by contrast, was
in a relatively small but unusually distinguished
laboratory in Copenhagen founded by her
parents, August and Marie Krogh (11).
Stanley G. Schultz

Stanley G. Schultz, University of Texas Medical

School, has been a distinguished leader in
physiological research and has provided outstanding
service to the profession of physiology in general
and to the American Physiological Society in
particular for many years. He has served as a
member of Council, as APS President, and as
chairman of the Long-Range Planning Committee.
He was instrumental in establishing the
Distinguished Lectureships of the APS sections. He
has also served as Editor-in-Chief of Physiological
Reviews, and as Editor-in-Chief of News in
Physiological Sciences. He served as the editor of the
2nd edition of the Handbook of Physiology – The
Gastrointestinal Tract.
John Severinghaus
was born in Madison, Wisconsin on May 6,
1922, under chloroform anesthesia requiring
version from a transverse lie and foot first
extraction. His Mother’s obstetrician
suspected the baby was brain damaged, and
was not relieved of that worry until he
attended the dinner at which the AOA
scholarly medical fraternity admitted
Severinghaus to membership after his first
year of medical school.
Novera Herbert Spector
Herb Spector writes: “In my ninety-first year, I
am looking more forward than backward. If our
species survives another 30 years, we will look
back in horror at the barbaric practices of
today in clinical medicine, just as we look back
in horror today at the methods used by well-
intentioned physicians in colonial times when
the standard treatment for most diseases was
phlebotomy. Using the best tools of the day,
they bled George Washington to death and
probably killed thousands of others. Today, too
many clinicians use whole body radiation or
chemotherapy to treat cancer, thus destroying
the body’s natural defenses and repressing
natural immunity so that it becomes a race to
see whether the cancer or the patient dies
first. Too often the patient loses the race.
Aubrey Taylor
June 4, 1933, El Paso, Texas
Aubrey Elmo Taylor was born in El Paso and grew up
in Bryson, Texas, Ventura and Oxnard, California, and
Fort Worth, where he was graduated from Paschal
High School, a school popularized by novelist Dan
Jenkins. After a hitch in the US Army and brief
employment with the meat processor Swift &
Company, Taylor enrolled at Texas Christian
University where in 1960 he received a baccalaureate
degree with a double major of mathematics and
psychology.
Charles M. Tipton (Tip) was born in Evanston, Illinois in
1927. Subsequently, his family moved to rural  Maryland
located half-way between Washington D.C. and Leesburg,
Virginia.  In 1945 he enlisted in the Army and was selected
to be a physical training instructor for stateside basic
trainees and for members of an infantry company in Japan.
He received a B.S. degree in Physical Education from
Springfield College in 1952 and an M.S. degree in the same
discipline a year later from the University of Illinois with
Professor Thomas K. Cureton as his advisor. After two years
of teaching general science, biology, physical education and
coaching four varsity sports in the high schools of Illinois, he
accepted an Assistantship  in Health Education to supervise
the exercise therapy for disabled students at the University
of Illinois in Urbana with Professor Howard Hoyman as his
Ph.D. advisor. Physiology was selected to be his minor.
Mario Vassalle
May 26, 1928, Tuscany, Italy
Mario Vassalle was born in Viareggio, a lovely
seaside resort on the Tyrrhenian coast of
Tuscany, not far from Pisa, Lucca and Florence,
where the golden sand and the azure of the
sea is crowned by the majesty of the nearby
mountains. There, he grew and received his
elementary and high school education. He
attended the Liceo Classico, which, in those
formative years of his youth, shaped his heart
and mind with long lasting effects: to this
day, he is deeply grateful for instilling in him
the love of beauty and a passion for clarity.
He then enrolled in the Faculty of Medicine at
the University of Pisa, where, once he obtained
his MD degree, he was an assistant in Medicine
for some 5 years.
Stephen F. Vatner

Is currently Professor and Director of the

Cardiovascular Research Institute at the University
of Medicine and Dentistry of New Jersey – New
Jersey Medical School. He received his B.A degree
from Grinnell College in Iowa and went on to
medical school at New York University.  Vatner did
his intern and residency training at the University
of Virginia and Postdoctoral Fellowship in the
Department of Physiology and Biophysics,
University of Washington, Seattle.  He has held
academic positions at the University of California,
San Diego, Harvard Medical School, Allegheny
University of the Health Sciences, and Weis Center
for Research, Penn State College of Medicine.  He
joined University of Medicine and Dentistry of New
Jersey – New Jersey Medical School in 2000.
Karlman Wasserman
Karlman Wasserman was born in Brooklyn,
New York in 1927 and graduated from high
School in Passaic, New Jersey in June
1944.  World War II was still quite active,
allied troops just having landed in
Normandy in their European campaign. 
Not having yet reached the age of 18, he
joined the U.S. Army in the Army
Specialized Training Reserve program to
start college education in basic engineering
at Princeton University.  After finishing
three quarters of college training at
Princeton, he was called to active duty in
April, 1945, having already reached the
age of 18 years
John B. West
John West was born in Adelaide, Australia in
1928. He had the good fortune to attend an
excellent high school and developed a love of
science, particularly high energy physics. He
considered this for a career but as was the
custom then (and still is) he moved straight from
high school to one of the faculties of the
University of Adelaide and, for various reasons,
chose medicine. He graduated with a medical
degree in 1951 after the six years' course at the
age of 23. After a year of residency he moved to
London, partly because academic medicine was
not well developed in Adelaide at the time, but
also because he wanted to see the world.
My 54 Years as a Physiologist 
Having enjoyed autobiographical reports from
the many very durable scientists you have been
publishing in The Physiologist, I am delighted to
get my turn, after I celebrated my 80th birthday
last week. Your invitation and questions
naturally evoked memories and reflections on
the shape of my whole career, so I begin with a
sketch of its six epochs, before presuming to
offer fragments of ( as you put it)  “wisdom to
pass on to (my)  younger colleagues”. I
composed this reply, in outline, as I was skiing
on a blue sky day, in good snow, at Park City
Mountain Resort in Utah. As a result, my
endorphins are high as I write.
October 17 al 21 Chicago
45th annual meeting, October 17-21, in Chicago
Exposición de Abstracts
Exhibit at the World's Largest
Neuroscience Marketplace
 ELECTROFISIOLOGIA
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• Sinápsis eléctrica
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adecuadas
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 COMPARTIMENTOS CORPORALES

• COMPARTIMENTO INTRACELULAR

• COMPARTIMENTO EXTRACELULAR:
- INTERSTICIAL
- PLASMA

CONTIENEN LIQUIDOS CORPORALES

 LÍQUIDOS CORPORALES:

Agua Corporal Total : 60%

LIC
40 %

Líquido Intersticial 15%

LEC

Plasma 5%

LIC = Liquido Intracelular

LEC = Liquido Intersticial + Plasma
 DISTRIBUCION Y VOLUMENES DEL AGUA EN
LOS COMPARTIMENTOS CORPORALES

COMPARTIMENTO % Litros
• LIQUIDO INTRACELULAR : 40 28.0
• LIQUIDO EXTRACELULAR
PLASMA : 5 3.5
INTERSTICIAL : 15 10.5
TOTAL : 60 42.0

Peso: 70 Kg. ACT: 60 %

INGESTIÓN Y PÉRDIDA NORMAL DE LÍQUIDOS EN UN ADULTO
QUE CONSUME 2,500 CALORIAS DIARIAS

Ingestión Eliminación

Cantidad Cantidad
Ruta Ruta
(ml) (ml)

• Agua en alimentos 1,000 • Piel 500

• Agua proveniente de • Pulmones 350
oxidación 300 • Heces 150
• Agua por ingestión oral 1,200 • Riñones 1,500

Total 2,500 Total 2,500

 COMPOSICIÓN DE LOS LÍQUIDOS CORPORALES

LEC (plasma + intersticial) LIC

Na+................................. 142 mEq/l Na+................................. 10 mEq/l
K+................................... 4 mEq/l K+.................................. 140 mEq/l
Ca+................................. 2.4 mEq/l Ca+.............................. 0.0001 mEq/l
Cl-.................................. 103 mEq/l Cl-................................... 4 mEq/l
HCO3-.............................. 28 mEq/l HCO3-............................... 10 mEq/l
Fosfatos.......................... 4 mEq/l Fosfatos............................ 75 mEq/l
Glucosa............................ 90 mg/dl Glucosa...........................0 a 20 mg/dl
Aminoácidos................... 30 mg/dl Aminoácidos.................. 200 mg/dl
Proteínas…………. 74 mEq/l

OSMOLALIDAD : 295 mosm/L 295 mosm/L

PH : 7.40 7.10
MEMBRANA CELULAR
 MEMBRANA CELULAR
• Relaciona el extracelular con el
intracelular .
• Medio de transporte
• Medio de comunicación
• Selectividad
• Es un sistema de relación
 MECANISMO DE TRANSPORTE DE LA
MENBRANA CELULAR

• TRANSPORTE PASIVO
1. DIFUSION
A) Osmosis
B) Gradiente Química
C) Gradiente Eléctrico
D) Gradiente Electro Químico
2. DIFUSION FACILITADA
A) Para Transporte de Sustrato
B) Para Transporte de Agua
3. FILTRACION
TRANSPORTE ACTIVO
1. Simporte (Unidireccional)
2. Antiporte (Intercambio) :Bombas
3. Exocitosis
4. Endocitosis
A) Fagocitosis
B) Pinocitosis
 OSMOSIS Y PRESIÓN OSMÓTICA

• Las
soluciones isotónicas hipertónicas
HIPERTÓNICAS contienen
mayor cantidad de solutos .
Las HIPOTÓNICAS contienen
menor cantidad de solutos.
Las soluciones ISOTÓNICAS
tienen concentraciones
equivalentes de solutos y, al
existir igual cantidad de hipotónicas
movimiento de agua hacia y
desde el exterior, el flujo neto
es nulo
NaCl 9 %0
Glucosa 5 - 20 %
TRANSPORTE ACTIVO
 GRADIENTE ELECTROQUIMICO.
• GRADIENTE QUÍMICO O DE CONCENTRACION:
- Es la diferencia de concentración de partículas entre
los compartimientos corporales.
- Causa un movimiento neto de iones del medio de
mayor al de menor concentración.
• GRADIENTE ELÉCTRICO:
- Es la diferencia de cargas iónicas entre dos
compartimientos
- Diferencia de potencial eléctrico (voltaje) a través de
la membrana celular. El citoplasma, es eléctricamente
negativo en relación con el líquido extracelular.
Equilibrio y potencial electroquímico.
ECUACIÓN DE NERNST.
• Una membrana separa una solución acuosa en
dos cámaras (A y B). El ión X+ se encuentra en
mayor concentración en el lado A.
• Si no existe una diferencia de potencial
eléctrico entre A y B, X+ tiende a difundir
desde A hacia B.
• Sin embargo, el lado A es eléctricamente
negativo. La tendencia de X+ de difundir de A a
B por la diferencia de concentración
permanece, pero ahora X+ tiende también a
desplazarse en el sentido opuesto (de B a A)
por la diferencia de potencial eléctrico a
través de la membrana.
• El sentido del desplazamiento neto de X+
depende de si es mayor el efecto de la
diferencia de concentración o el del potencial
eléctrico:
Fuerza de concentración VS Fuerza eléctrica
 Fuerza Eléctrica (Fe)
• Esta relacionada directamente a la diferencia de
potencial a ambos lados de la membrana
Fe = zF (E ab)

Z = carga del ion, +1 o -1 para Na, K, Cl.

F = constante de faraday 96,500 culb/mol
E = diferencia de potencial a ambos lados de la
membrana. Si E es expresado en voltios, la Fe
esta dada en joules/mol
 FUERZA DE CONCENTRACION
• No esta relacionada al gradiente de concentración si
no a la proporción logarítmica de las
concentraciones.
Fc = RT ln C2/C1
R= constante de los gases 8,31 J/mol
T= grados celcius + 273 = 310 ºk

Multiplicando Rx T da resultado en Joules/mol, lo mismo

que en la ecuación de la fuerza eléctrica, entonces podemos
sumar o restar ambas ecuaciones directamente.
 Diferencia de potencial electroquímico.
Si llamamos µ1 y µ2 al potencial electroquímico de los compartimientos 1 y 2
respectivamente, este será:
µ1 = RTlnC1 + zFV1
µ2 = RTlnC2 + zFV2

La diferencia de potencial electroquímico será:

∆µ = µ1 - µ2 = RTln C1/C2 + zF (V1 – V2)

Si para que haya equilibrio, no debe haber diferencia de potencial electroquímico,

se debe cumplir que:
µ1 - µ2 = 0 entonces: IGUALANDO LAS ECUACIONES DE Fc y Fe

RTln C1/C2 = - zF (V1 – V2)

(V1 – V2) = ∆V = RT/-zF ln C1/C2 , ∆V = diferencia de potencial eléctrico.

V1 – V2 = RT/zF ln C2/C1 ECUACIÓN DE NERNST.

 ECUACIÓN DE NERNST:

DV = RT. ln C2
zF C1

DV = diferencia de potencial eléctrico

potencial de equilibrio (E)
R= 8,31 J/mol.
T= 310 ºK
F= 96,500 cb/mol
76
 ECUACIÓN DE Nernst
• Define el potencial de equilibrio de un ion(E).
• Se dice que un ion esta en equilibrio, cuando las
fuerzas que la mueven (fuerza de concentración
y eléctrica) son iguales y opuestos, de tal manera
que no hay un movimiento neto a través de la
membrana.
• Permite calcular individualmente el potencial de
equilibrio de una especie iónica a partir de sus
concentraciones a ambos lados de la membrana.
 Ecuación de Nernst.
• Muchas veces conviene convertir la ecuación de Nernst a una
fórmula que implique logaritmo en base 10.
lnx = 2,303 logx.

• Debido a que los potenciales biológicos se expresan en

milivoltios (mV), hay que seleccionar las unidades de R de
forma que RT/F se expresen en milivoltios. La cantidad
2,303.RT/F es igual a 60 mV, valor que se mantiene
aproximadamente para la mayoría de las condiciones
experimentales en biología, y una forma útil de emplear la
ecuación de Nernst es:
V1 – V2 = 60 mV/z log C2/C1
 La Ecuación de Nernst
RT [ X ]out
potencial eléctrico EX  ln concentración extracelular
del ion X zF [ X ]in concentración intracelular
valencia
del ion X

RT/F es constante ( 0.061 a 38° C)

79
Potencial de Equilibrio de Nernst para el K+

Calcular el potencial de equilibrio de Nernst para K+:

1. Asumir que [K+]i es 10 veces mayor que [K+]o
2. [K+]o/[K+]i = 0.1
3. Valencia = +1

0.058 [K[ K]o]o

EKK 00.058
.058 log
(log
log(1)0.1-58
)  mV
-0.058V
z [K[ K]i]i

80
Potencial de Equilibrio de Nernst para el Na+

Calcular el potencial de equilibrio de Nernst para Na+:

1. Asumir que [Na+]o es 10 veces mayor que [Na+]i
2. [Na+]o/[Na+]i = 10
3. Valencia = +1

0.058 [ Na
[ Na]o]o
Na 00
ENa .058
.058 log
 (log(
log ) 0.058V
1)10  mV
58
z [ Na
[ Na]i]i

81
Potencial de Equilibrio de Nernst para el Cl -

Calcular el potencial de equilibrio de Nernst para Cl-:

1. Asumir que [Cl-]o es 10 veces mayor que [Cl-]i
2. [Cl-]o/[Cl-]i = 10
3. Valencia = -1

0.058 [Cl[Cl
]o ]o
EClCl 00.058
E .058 log
log
1log( ) mV
-58
-0.058V
10
z [Cl[Cl
]i ]i

82
 Potenciales de Equilibrio de ..

• E ion Cl = - 70 mV.
• E ion Na = + 60 mV (toda la despolarización)
• E ion K = - 90 mV(cerca al potencial de reposo
 EQUILIBRIO DONNAN
[Na+]2 = [K+]2 = [Cl-]1
[Na+]1 = [K+]1 = [Cl-]2 INTRACELULAR EXTRACELULAR
mEq/L meq/L

En el EQUILIBRIO, la
Na+ 10 142
relación entre iones de
ambos lados de la membrana K+ 140 5
es un valor constante.
Cl- 7 103
La célula NO ES UN SISTEMA EN
EQUILIBRIO. Prot- Muy elevada
165 mM

85
 ECUACIÓN DE
GOLDMAN-HODGKIN-KATZ
• Si la célula fuera un sistema en equilibrio no debería
haber flujo neto transmembrana.

• La célula es un sistema en ESTADO ESTACIONARIO en

donde hay un flujo neto que siempre se mantiene
constante por determinados mecanismos
Homeostásicos (se usa la ecuación de Goldman…….)

86
Ecuación de Goldman ó de Goldman-Hodgkin-Katz

• Calcula el potencial de equilibrio cuando mas de un ion

es permeable.
• Incorpora los coeficientes de permeabilidad de cada ion
(especifico de cada membrana)

RT PK [ K  ]o  PNa [ Na  ]o  PCl [Cl  ]i

Eions  ln   

F PK [ K ]i  PNa [ Na ]i  PCl [Cl ]o

87
 POTENCIAL DE MEMBRANA EN REPOSO
(PMR)
• Interior de las células hay mas ANIONES que CATIONES
• Hay mayor cantidad de cargas negativas adentro.
• Se crea una diferencia de potencial eléctrico entre
ambos lados de la membrana en reposo llamado
POTENCIAL DE MEMBRANA EN REPOSO
• Se escribe con signo (-) significando que en el interior es
negativo con respecto al exterior.
• Su valor varia según tejido : – 10 mV en GR hasta -90mV
en fibra muscular estriada.

88
 FENÓMENOS QUE DETERMINAN EL
POTENCIAL DE MEMBRANA.

• Bomba ATPasa Na-K (3Na-2K) genera negatividad adicional

(5 a 20%).
• Efecto Donnan: distribución asimétrica de los iones por
la no difusión de aniones proteicos.
• La casi nula permeabilidad de la membrana para los
aniones proteicos.
• La baja permeabilidad de la membrana celular en
reposo a los iones de Na.
• La relativamente elevada conductancia de la
membrana celular en reposo para el K.
• 20 a 100 veces más permeable al K+ Gradientes de
concentración para Na+ y K+.
 Cambios en el
potencial de membrana
Potencial local (electrotónico)
- Variable
- Pasivo
- No se propaga (se extingue rápidamente)

Potencial de acción
– Siempre igual (“todo o nada”)
– Activo
– Se propaga sin cambios
 POTENCIAL LOCAL (ELECTROTÓNICO)

Umbral 1 ms
POTENCIAL ELÉCTRICO

0 mV

-70 mV

TIEMPO
POTENCIAL DE ACCIÓN

92
Los Potenciales Lentos disminuyen a medida que se desplazan

 Si un estímulo no cambia el
potencial de membrana
llevándolo hacia valores
positivos, la señal muere y la
neurona no responde
disparando un PA.
 La cantidad de cambio del
potencial de membrana
necesario para generar un
potencial de acción es el
potencial umbral.
 POTENCIAL DE ACCIÓN
• La existencia de un PMR es la base de la
excitabilidad de las células (responder a
ciertos estímulos)
• Cuando una célula recibe un estímulo, su
membrana sufre una serie de cambios que
modifican el PMR.
• Dichos cambios se conocen como POTENCIAL
DE ACCIÓN.

94
 MECANISMO FÍSICO DEL P.A.
• “El potencial de acción es consecuencia de los
cambios de permeabilidad a ciertos iones”.
• ETAPAS
– Despolarización.
– Repolarización.
– Hiperpolarización.

95
 POTENCIAL DE ACCIÓN

Potencial de acción: cambio

rápido en el PMR y retorno a
la situación inicial

PA permite en células
excitables: transportar
señales
• Después la membrana se vuelve a hacer
impermeable al Na+ y se restablece el potencial
de reposo.
 Potencial de acción
+50
mV
umbral
0
Depolarización Repolarización
-90 mV hacia 0 mV (0 mV hacia -90 mV)
-50
Hiperpolarización
(potencial se vuelve más negativo que PMR)

-100
0 1 2 msec
REGISTRO y PARTES DEL POTENCIAL DE ACCIÓN
 POTENCIAL DE ACCIÓN

1. Potencial de membrana en reposo

2. Estimulo depolarizante umbral:
apertura canales Na+ Voltaje-
Dependientes
3. Entrada rápida de Na+:
depolarización
4. Cierre canales Na+, apertura Canales
K+
5. Salida de iones K+: hiperpolarización
6. Canales de K+ siguen abiertos, iones
K+ siguen saliendo (periodo
refractario absoluto y relativo)
7. Vuelta a potencial en reposo

POTENCIAL DE MEMBRANA
101
 Bases iónicas del potencial de acción

Los PA son causados por la apertura de canales para Na+ y K+

– REPOSO: cerrado, pero

disponible para su
apertura por estímulos
químicos o eléctricos.
– ACTIVO: abierto,
permite el paso de una
corriente iónica.
– INACTIVO: cerrado, y
NO disponible para su
apertura
 CONDUCTANCIA PARA EL SODIO-VOLTAJE DEPENDIENTE = PA

APERTURA DE CANALES DE SODIO

UMBRAL - 40mV APROX. ENTRADA MASIVA DE SODIO A CÉLULA

+35mV
DESPOLARIZACIÓN
0mV

SE ABREN MÁS CANALES DE SODIO

EL POTENCIAL TIENDE AL PUNTO DE
EQUILIBRIO DEL Na+

¿Cómo termina este

feed-back positivo?
-40mV
Por inactivación de la
-
Tiempo en ms
conductancia para el
60mV Conductancia Na+ ConductanciaK+ Na+
 CAMBIO EN LA Disminuye la
ESTÍMULO Ingresa Na +
Carga (-) adentro.
PERMEABILIDAD
AL Na+ DESPOLARIZACIÓN

La permeabilidad al K+
Comienza a salir K+ Inmediatamente que
cambia tanto que sigue
y se restablece la disminuye la
saliendo K+ haciendo
carga negativa permeabilidad al
mas negativa de lo
adentro. Na+, aumenta al K+.
normal.
REPOLARIZACIÓN
HIPERPOLARIZACIÓN

ADENTRO

La bomba de Na+ y K+
regenera el valor de
PMR.

AFUERA
104
 MECANISMO MOLECULAR DEL P.A.
¿PORQUÉ CAMBIA LA PERMEABILIDAD DE LOS IONES Na+
Y K+ CUANDO SE GENERA EL POTENCIAL DE ACCIÓN?

Cambia la
Cambia la
permeabilidad
DESPOLARIZACIÓN permeabilidad
ESTÍMULO al Na+.
al Na+.
Se abren
Se cierran
canales de Na+
canales de Na+

Los canales Cambia la

HIPERPOLARIZACIÓN de K+ siguen REPOLARIZACIÓN permeabilidad
abiertos al K+.
Se abren
canales de K+
Bomba de Na+ y K+ PMR

105
PROPAGACIÓN DEL POTENCIAL
DE ACCIÓN

106
PODEMOS MEDIR EL NUMERO DE POTENCIALES QUE PASAN POR UNA MEMBRANA EN EL TIEMPO Y
RELACIONARLO CON LA INTESIDAD DEL ESTIMULO .
Conducción del potencial de acción

 Los PA y respuestas subumbrales

se propagan por flujos de corriente
locales

 El mecanismo de conducción es
conocido como conducción
electrotónica.
 Se abren canales de Na+
dep. de voltaje y
el Na + entra en la cel

Se produce un potencial
por encima de umbral

La entrada de Na+
despolariza la membrana que
abre más canales de
Na +

Las cargas positivas fluyen a zonas

adyacentes de la membrana por
flujos de corriente local
 Región Región Región
refractaria activa inactiva

La zona despolarizada primero está en En las partes distales, la corriente local

período refractario. Los canales de K+ se de la región activa causa despolarización
han abierto y los de Na están cerrados. Sale de nuevas zonas de la membrana.
K+ del citoplasma, se repolariza la
membrana
 Propagación del potencial de acción
El PA conduce el impulso sin decremento para ello el PA se regenera a lo largo de la fibra y se
dice que es propagado además de conducido.

Cuando un área del axón

alcanza el umbral, el influjo
de Na+ y la generación del
PA se repetirá una y otra vez
en una dirección en cada
segmento de la membrana a
lo largo de la célula
excitable.

El tamaño y la forma del PA permanece invariable, sólo se permiten variaciones en la

frecuencia de disparo para transmitir señales a lo largo de la fibra. La máx. frecuencia está
limitada por la duración del período refractario absoluto(≈1 msec) a aprox. 1000 impulses/sec
en nervios grandes.
 Velocidad de conducción

Variables de las que depende la velocidad de conducción:

• Diámetro de la fibraFibras más grandes en diámetro tienen
velocidades mayores. Esto es debido a un descenso en la
resistencia a la conducción según aumenta el radio ( del que
depende el área de sección)

• Grado de mielinización
Mielinizadas mayor velocidad
 Efecto de la mielinización en la velocidad de conducción

Mielina vueltas de la membrana

plasmática de las células deI Schwann que
se enrrollan alrededor de los axones
nerviosos ( mas de 100 capas de membrana
plasmática) .

 Nodos de Ranvier: interrupciones cada 1-

2mm, se corresponden con los espacios entre
dos cél de schwann.
Velocidad de conducción

Na+

Na+

mielina

Na+
++++++++++++++++++++++++++++
A la membrana llega un estímulo.

---------------------------

++++++++++++- -+++++++++++++ Provoca la salida de K+ y el

ingreso de Na+.

-- - - - - - - - - - - ++ - - - - - - - - - - - - - Si el estímulo es de intensidad
suficiente, se llega al valor
umbral y se produce el Potencial
++++++++ + - - - - - - - - +++++++++ de Acción que se propaga en
todas direcciones
- - - - - - - - - ++++++++ + - - - - - - - - -

Si el estímulo NO es de
++++++++++++++++++++++++++++ intensidad suficiente, las fuerzas
regeneradores RESTITUYEN el
PMR y no se logra la generación
--------------------------- de un PA propagable.
PMR

Este tipo de conducción del PA se realiza en las

neuronas de tipo AMIELINICAS.
116
CONDUCCIÓN SALTATORIA

Este tipo de conducción del PA se realiza en las

neuronas de tipo MIELINICAS (mas veloces).
117
PROPIEDADES DEL POTENCIAL DE
ACCIÓN

118
• UMBRAL DE EXCITACIÓN: para que se genere
el PA, la intensidad del estímulo ha de ser tal
que supere cierto valor de potencial (20 mV
para la neurona)
• LEY DEL TODO O NADA: un estímulo
supraumbral origina un PA que una vez
producido se transmite a lo largo de toda la
neurona. Si el estímulo es subumbral, el PA no
se produce.
119
• PERÍODO REFRACTARIO: después que se ha producido el PA,
existe un tiempo en el cual la célula no responde a los
estímulos, posteriormente la célula va recuperando su
excitabilidad: Periodos Refractarios Absoluto y Relativo.

• VELOCIDAD DE PROPAGACIÓN: la velocidad con la que se

conduce un impulso nervioso es constante. Depende de:
– Diámetro de la fibra.
– Presencia o ausencia de mielina.
– Propiedades eléctricas de la membrana: resistencia,
capacitancia, etc.
120
 Período refractario

Período durante el cual es imposible generar

otro potencial de acción y coincide con la primera parte del PA

Absoluto Relativo Un gran número de canales de Na+ son

inactivados y no pueden volver a abrirse
hasta que la membrana se repolariza

Período refractario relativo. Durante la

última parte del potencial de acción la cél
es capaz de disparar un nuevo potencial
pero se necesita un estímulo mayor de lo
normal. La conductancia al K+ está
aumentada.
PROPIEDADES ELECTRICAS
K+ (5), Na+ Mg++ 2,5, Cl- (103), Ca++ (5)
(140),
+ + + + + + + + + + + + ++++ Reposo
+ + 0
+ --------------- +
+ - A-, K+(150), Na (10), - + + -90 mV
+ - Mg (40) ++
- +
+ --------------- +
+++++++++++++++ Célula polarizada

- - - - - - - - - ++++++
E
- + +
s
+++++++ - - - - 0
t
- +
i
+ K -
m
- + -90 mV
u
+ Proteínas -
l
- +
o
+++++++- - - -
- +
- - - - - - - - - ++++++ Despolarización
 Repolarización
+ + + + + + Na - - - - - -
PAT
+
+ ----------++++ - 0
+ - K + -
+ - - -90 mV
Proteínas +
+ ----------++++ -
+ -
+++++++++- - - - - -

+ + + + + + + + + + + + ++++
+ - + 0
--------------
+ +
- A-, K+(150), Na (10), -
+
-90 mV
+ +
- Mg (40)
++
-
+ +
---------------
+ +
+++++++++++++++ Célula polarizada
 CONDUCCIÓN NERVIOSA
• Un axón logra conducir en cualquier dirección in vitro.
• En el animal vivo, los impulsos se propagan en un solo sentido,
ya que, tras el paso de un PA, cada segmento de la fibra pasa
por una fase refractaria, durante la cual no puede ser excitada.

• La conducción en un solo sentido se llama ortodrómica

• La conducción en la dirección opuesta se llama antidrómica

• La velocidad de conducción depende también del diámetro de

la fibra nerviosa; cuanto mayor es el diámetro, y por tanto
mayor el área de sección transversal, menor es la resistencia
longitudinal del axón, mayor es la velocidad.
 TRANSPORTE AXOPLASMICO

• Transporte a través de axon de sustancias del soma hasta lo

botones terminales y viciversa.
• Bloqueadores: Anoxia, acidez, cianuro, N2

• TRANSPORTE ANTEROGRADO (Ortodrómico)

Las sustancias se transportan de soma a la terminación
nerviosa
– Transporte rápido: 40 mm/día (3 mm/hora), vesícula
sináptica, depende del ATP.
– Transporte lento (6-10 mm/día),
Transporta elementos del Cito esqueleto
– Transporte muy lento (0.5 – 3mm/día)
Transporta proteínas del Cito esqueleto y tubulina
 TRANSPORTE AXOPLASMICO

TRANSPORTE RETROGRADO (antidromico)

• Las sustancias se transportan desde los
terminales al soma.
• Velocidad 200 mm /día
• Factor de crecimiento nervioso, algunas
vesículas, virus, toxina tetanica, peroxidasa
del rabano picante