ALFRED BENNUN, PH.D.

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SCHOLARLY AND CREATIVE

TEACHING

1957 - Research technology, non-credit course, Dept. of histology and Embryology, School of Medicine, Univ. of Cordoba, Argentina

1958 - Sections of Experimental Biochemistry, Dept. of Biochemistry, School of Biochemistry, University of Buenos Aires

1958 – 1962 - Sections of Biological Chemistry, School of Medicine, University of Buenos Aires

1967 - 1969 - Biology 103-104, General Biology, Dept. of Biology, University of Puerto Rico

1967 - 1969 - Biology 379, Reading and Research, Dept. of Biology, University of Puerto Rico

1968 - 1969 - Biology 354, Cellular Biochemistry, Dept. of Biology University of Puerto Rico

1968 - 1969 - Biology 257, Plant Physiology, Dept. of Biology, University of Puerto Rico

 

990:494 Seminar in Zoology (1 cr)
Fall 1970, Spring 1976, Spring 1977.

115:413 Experimental Biochemistry (2 cr)
Fall 1971, Fall 1972, Fall 1973, Fall 1975, Fall 1980.

990:411 Endocrinology
Fall 1977.

115:414 Experimental Biochemistry (2cr)
Spring 1972, Spring 1973, Spring 1974.

990:204 Physiological Zoology (4 cr)
Spring 1976, Spring 1977, Spring 1978

120:101 General Biology Laboratory
Fall 1973, Spring 1974, Fall 1986

115:310 Elementary Biochemistry
Spring 1980, Spring 1984, Spring 1985, Spring 1986.

115:312 Elementary Biochemistry
Fall 1980, Fall 1983, Fall 1984, Fall 1986.

115:404 General Biochemistry
Spring 1980, Spring 1981, Spring 183, Spring 1984, Fall 1984, Spring 1985.

760:407 General Physiology
Fall 1981, Fall 1982, Spring 1979, Spring 1983.

115:405 Problems in Biochemistry
Fall 1982, Fall 1983, Spring 1984, Spring 1985.

990:491 Special Problems in Zoology
Fall 1982, Spring 1983.

 

Courses Taught (at Graduate School Rutgers University)

115:513 Metabolic Pathways Controls (3 cr)
Fall 1969, Fall 1970, Fall 1971, Fall 1972, Fall 1973, Fall 1975.

990:513 Metabolic Pathways Controls (3 cr)
Fall 1975, Fall 1976, Fall 1977, Fall 1979.

115:514 Metabolic Pathways Mechanisms (3 cr)
Spring 1970, Spring 1971, Spring 1972

990:514 Metabolic Pathways Mechanisms (3 cr)
Spring 1973, Spring 1974, Spring 1976, Spring 1977, Spring 1978.

990:511 Advanced Endocrinology (3 cr)
Fall 1976, Fall 1977, Fall 1979, Fall 1980, Fall 1981.

990:616 Special Topics in Biochemistry (1 cr)
Fall 1975, Spring 1976, Spring 1977, Fall 1977, Spring 1978, Spring 1979, Spring 1984, Fall 1984, Spring 1985, Fall 1985.

115:606 Advanced Studies in Biochemistry
Spring 1974

115:701 Research in Zoology
1969 to Present

990:509 Advanced Problems in Zoology
Summer 1976, Spring 1977, Summer 1978, Spring 1979, Fall 1978, Fall 1985.

160:572 Biochemistry
Spring 1980, Spring 1981, Spring 1984, Spring 1985, Spring 1986

 


Supervision of Honors Thesis Completed by Students in the Undergraduate Honors Program:

1969 1968 -        W. Velez-Vega and M. Dieppa - Honors Program Advisor -Biology Department University of Puerto Rico

1981 - Nancee A. Novak - The effect of cyclic nucleotides upon glycolysis in the human erythrocyte

1982 - Bessie B. Fouces - Kinetic studies on brain adenylate cyclase

1984 - Sylvia Menendez - The role of energy charge in the regulation of brain adenylate cyclase activity

 

Summer Supervision of Minority High School Students with Research Apprentice Program of NIH-BRSP

1982  Patricia Grady

1983  Patricia Grady

1984  James Anderson

1986  Manuel Dasilva

 

VII. UNIVERSITY SERVICE

1967 – 1969 - Member of the Graduate Faculty, Department of Biology, University of Puerto Rico

1970 - Full member of the graduate School of Rutgers University.

1970 - Full member of the cooperative Graduate Program in Biochemistry, Rutgers University.

1970 – 1973 - Member of the University Science Council, Rutgers University

1975 - . Member of the Academic Standards Committee of the Cooperative Graduate Program of Biochemistry.

1970 - Member of several graduate students’ committees in the Graduate Program in Biochemistry

1970 - Full member Graduate Program in  Zoology. NCAS

1970 - Member of several graduate students  committees in the Graduate Program in Zoology NCAS

1975 – 1978 - Member of the NCAS Academic Affairs Committee

1977 – 1978 - Member of NCAS Graduate School Planning Committee

1979 – 1981 - Member of NCAS Graduate School Planning Committee

1977 – 1978 - Member of Curriculum Committee, Dept. of Zoology and Physiology NCAS

1978 - Member of Executive Committee of the Graduate Program in Zoology NCAS

1979 - Member of Teaching Evaluation Committee NCAS

1978 - Member , Cancer Coordinating Committee of Rutgers, Newark NCAS

1983, 1985 - Member, Executive Committee Graduate Program in Zoology NCAS

1984 – 1986 - Member of the advisory committee for MBRS program in Newark Campus of Rutgers.

 

 

VIII. GRADUATE STUDENT SUPERVISION

Supervision of Ph.D. candidates research and titles of Ph.D. dissertations completed by students under my direction and initial post-doctoral positions:

 

Susan B. Golz - Regulation of brain adenylate cyclase by physiological and pharmacological factors.  Ph.D. in Zoology awarded May 1976. Initial Position: Assistant Professor, Bergen Community college.

Robert H. Harris - Hormonal regulation of rat fat cell CAMP levels and fat ceil membrane adenylate cyclase.  Ph.D. in Biochemistry awarded November 1977.  Initial Position: Research specialist, Division of Metabolism of Food and Drug Administration, Bethesda, MD.

Heripsime Ohanian - The role of divalent metals, adenosine 5'-triphosphate and hormones in the regulation of rat brain adenylate cyclase.  Ph.D. in Zoology awarded June 1980.  Initial Position: Medical School, Lille University, France.

Vincent A. DeBari - Regulation of the hexose monophosphate pathway in cyclic nucleotides and divalent cations in the human erythrocyte. Ph.D. in Zoology awarded May 1981. Initial Position: Director, Renal laboratory, St. Joseph's Hospital, Paterson, N.J.

Joseph W. Sulner - Renal renin release - a stimulatory role for Calcium Ph.D. in Zoology awarded May 1984.  Postdoctoral Hoffman – Laroche. Initial Position: Associate Investigator N.Y. Veteran's Hospital.

Rafaela Cruz - The integration of hormones, Ng 2+ Ca2+ and their nucleotide complexes on the regulation of brain adenylate cyclase. Ph.D. in Zoology awarded October 1986. Initial Position: Adjunct Faculty in Anatomy and Physiology at Upsala College, East Orange, N.J..

Pasquale Vicario – The regulation of Insulin Receptor Tyrosine Kinase by Insulin, Divalent metal cations and metal – ATP substrate. Ph. D. in Zoology awarded May 1988. Initial Position: Investigator at Merck Company.

Christopher Casciano – The pharmacological characterization of the bicarbonate, proton, and mucus transport systems of the rodent gastric mucosa and their significance in cytoprotection. Ph. D. in Zoology awarded September 1988. Initial Position: Investigator at Schering Company.

 

Supervision of Ph.D. candidates research under my direction:

1984  G. Serban

1985  P. Uychich

1985  L. Rumennick

Supervision of Masters candidates research and Masters Thesis completed by students:

1975 - Chairman of the committee for the Master in Zoology of Susan Brydon Golz. M. Sc. awarded 1975.

1978 - Chairman of the committee for the Master in Zoology of Heripsime Ohanian. M. Sc. awarded 1978.

1981 - Chairman of the committee for the Master in Zoology of Rafaela Cruz. M. Sc. awarded 1981

Supervision of research:

1968 – 1969 - Advisor in the Biology Program to 1. Bulla, M.S. candidate at university of Puerto Rico

1970 – 1971 - Advisor in the Graduate Program of Biochemistry to E. Albanese and V. Cannistraro.

1970 – 1971 - Advisor in the Master in Zoology Sciences Program to E. Wallendjack

1977 - Research advisor to Kamran Borhanian, student at NJCMD, Rutgers Medical School

1977 - Postdoctoral research advisor to Salette de Farias on a leave from Institute Butantan, Sao Paulo, Brazil

1984 - Research advisor to Norbert Seidler, student at NJCMD, Rutgers Medical School

Membership on Graduate Student Committees:

1974 - Vincent Rama (Ph.D., Zoology)

1977 - Jeff Stevens (M.S., Biochemistry)

1980 - Jeff Stevens (Ph.D., Biochemistry)

1979 - Rosy Jordan (M.S., Zoology)

1979 - Gerald Costa (M.S., Zoology)

1980 - Robert Jonathan Weiss (M.S., Zoology)

1980 - Beryl Joyce Carby (M.S., Zoology)

1980 - Vera Minak (M.S., Zoology)

1980 - Frank R. Settineri (M.S., Zoology)

1981 - Diane G. Verga (M.S., Zoology)

1983 - Dolores Curtis (M.S., Zoology)

1983 - Marlene Blanco (M.S., Zoology)

1983 - Mark Schachman (M.S., Zoology)

1983 - Thomas Brankner (M.S., Zoology)

1984 - Lee D. Carpe (Ph.D., Zoology)

1984 - Dennis Guiliani (Ph.D., Zoology)

1985 - Eva-Pia Reich (Ph.D. Zoology)

1985 - Ilda D. D'Onofrio (Ph.D. Zoology)

1988 - John C. Anthes (Ph. D. Zoology)

 

IX. RESEARCH ACCOMPLISHMENTS

A. Active Transport and metabolic control

a)       It was demonstrated that active transport of phosphate by yeast is directly dependent on high levels of energy – rich phosphates maintained by either endogenous oxidation or by exogenous substrates during oxidation by either glycolysis or respiration.

b)      It was shown that yeast’s uptale of acetate, pyruvate, etc. and their oxidation rate within the Krebs cycle, rather than under direct control by oxygen itself, is subject to an energy dependent regulatory mechanism supported by ATP levels and their equilibria with other energy rich phosphate compounds.

 


B. Mechanism of oxidative phosporylation

         In 1963, the DNP – activated, Mg – dependent ATPase of yeast was for the first time isolated and characterized for its role in oxidative phosporylation.

III. Mechanisms of enzymatic induction and reversion as cells adaptative response to environmental changes.

         It was found that Wuglena controls endogenous phosphatase levels by a low phosphate contration – dependent induction of the synthesis of a phosphate. Induced and constitutive phosphataase were isolated and characterized as differente enzymes. It was found that disappearance of the induced enzyme (reversion) is a high phosphate shows a temperature – dependent function. The low activation energy (heat) required for the denaturation of the induced enzyme in solution allows to postulate that cell’s reversion operates through a high phosphate – dependent thermic sensitization of the induced enzyme which accelerates its thermic decay.

 

C. Mechanism of photophosphorylation

a)     In 1964, the proposition that high energy intermediates should be equated with conformational changes was published. It was proposed that conformational change  participated in energy conservation and transduction. Ca2+ and cysteine wiche eleicit light – requiring ATPases were shown to, modify the kinetic behavior of the ATP synthetase. ADP and GDP show inhibitory constants (Ki) for the light – requiring ATPases of the same numerical value that their Kms for residual and modified phtophosphorylation. This would suggest the participation of a single, but modigiable, active center in the mechanism which would determine the direction of the ATP synthetase activity

b)    The first successful resolution and reconstitution of an energy – dependent ATPase function was achieved for chloroplast light – requiring ATPases. Isolation and purification of the ATP-synthetase choroplast’s coupling factor. It was also shown that the latter becomes an ATPase by heat or trypsin treatmen.

c)     Reso1ution and reconstitution of the functlon for metal-dependent binding of the coupllng factor (ATP synthetase-ATPase) by the chloroplast membrane. Characterization of allotopic properties of the system.  Characterization of the Inhibitory capacity by the membrane on the ATPase activity during binding.

d)    The finding of the capability of the ATPase to maintain cooperative interaction with 2 ATP, 4PO4, 2 ADP and 16 molecules of water, was used to postulate that the active form of the ATPase is a structure characterized by conserving the energy of 14 H-bonds or 14 Ki1oca1ories (1 kilocaloráe per each solvating water molecule), where as that of the inhibited form of the ATPase is a estructure characterized by a decrease in energy content of the same value. This appears to indicate the potential of the ATPase itself to function as a high-energy intermediate during the turnover of the ATP synthetase system.

e)     First report on the allosteric kinetics of the ATPase (activated coupllng factor), and demonstration that proton concentration has the capability to requlate the modification of the ATPase from a form of high affinity for ATP and low affinity for phosphate and ADP, into another form of opposite properties.

 

D. Energy Transduction

 

a)     Kinetic and thermodynamic parameters were described for the operatien of ATPase coupled for ionic transport, muscle contraction, interaction between cytochromes and coupling factor (ATPase) and the effect of ions in the modification of chloroplast’s light-dependent activities.

b)    The postulation of energy transduction mechanisms resulting from the exergonic association of proteins with divalent metals (Me2+). The turnover of the protein te a dissociated state (endergonic) is driven by the chelation of the binding Me2+ by ATP4- (exergonic). ATPase activiy turns ATP4- into the waker chelating agent ADP3- allowing Me2+ dissociation and the return te the initial high concentratien of free Me2+.

c)     The binding of a divalent meta1 requirement for the EDTA-treated membranes binding of the coupling factor-ATpase led to the postulation that electron transfer is coupled to phosphorylation by cytochreine-dependent or pH-dependent modificatien of the meta1 chelated states of R-groups located in the coupling site of the ATPase.

d)    Consequently, the unitary hypothesis can be differentiated from the chemical hypothesis in that it postulates the formation of high energy intermediates by the transfer of groups across coordinative bonds rather than across covalent bonds. It also integrates conformational and chemiosmotic prepositions within a single framework.  Accordingly, the hypothesis uses a single thermodynamic and kinetic framework for findings that were previously explained only through the alternate use of three different hypothesis.

 

E. Hemoglobin

It has been previously proposed that in an open thermodynamic system, the turnover of the state of Me2+ as ligand of pretein(s) would yield the energy of activation for driving conformational and functional changes of protein.  Accordingly, it was proposed that oxygenation of Hb would lead Me2+ induce-fit a chelating configuration of side chains. This binding of Me2+ to Hb would dislocate proxima1 histidines from the positions reguired for steric hindrance and would allow Bohr side chanin(s) te release their proton(s).  Mg2+ rather than Zn2+, was suggested as the main physiological Me2+ to compete with Zn2+ at cellular concentrations of 11 ug/g red cells for the chelating side-chains of Hb. Oxygenation induces binding of Mg2+ or Zn2+   to Hb. Hence, according to the mode1 the PH-dependent stability order of the chelated complexes during deoxygenation would be: (Zn2+)2-oxyHb>(Zn2+, Mg2+)-oxyHb>(Mg2+)2-oxyHb

 

F. Characterization of the hormonal and metabolic parameters that control adenylate cyclase activation

 

a)     GH potentiates the effect of epinephrine, glucagon and theophylline increasing endogeneus cAMP levels in fat celis.

b)    Isolated membranes show that epinephrine activates adenylate cyclase by decreasing its Km for ATP and its Km for Mg2+ to reach a maximal activatory effect. Adenylate cyclase can be described as a receptor-coupled enzyme system in where the hormone binding by the receptor increases the affinity of the enzyme's regulatory site for Mg2+. The regulatory site also functions in the absence of epinephrine because high Mg2+ concentration is also capable of activating adenylate cyclase (19)

 

G. Characterization of hormonal and metabolic parameters that desensitize and inactivate adenylate cyclase

 

a)     It was found that an adenylate cyclase membrane preparation from rat's cerebral cortex, during prolongued preincunation at 37 C, became rapidly inactivated when noradrenaline was present.  Total loss of activity was reached at 100 minutes.  ATP was able to protect the enzyme against this effect for over 4 hours.  On the other hand, ATP alone activated the enzyme but desensitized it to its stimulation by noradrenaline.  The characterization of the described effect indicates a noradrenaline-dependent thermic sensitization which accelerates thermic decay. ATP protects against this effect.  Hence, It was surmised that since anxiety and stress are mediated by catecholamines, prolongued maintenance of high noradrenaline levels in conditions that exhaust endogeneus ATP in the affected organs, may inactivate adenylate cyclase and explain, therefore, the aetiology of psychosomatic and stress related diseases  (i.e. anxiety leading to depression).

b)    Solving multiple-equilibirum equations demonstrated that the relationship between energy charge (ATP vs. ADP and AMP) and relative concentration of free vs. chelated by nucleotides divalent metals (Me: Mg2+, Mn2+, Ca2+) plas a modulatory role for adenylate cyclase and insulin-dependent tyrosine kinase at three sites (substrate site, Me2+-dependet regulatory site and Me-requiring site for control of hormone responsiveness)

 

 

H. Metabolic controls

The cellular uptake of glucose and/or the lysis of glycogen it is expected to change the concentrations of chelating metabolites (sugar phosphates) and therefore the relative concentrations of free versus chelated divalent metals.  Thus a decrease in chelating metabolites constitutes an amplification mechanism of the adenylate cyclase (AC) response, since it simultaneously increases the concentrations of the substrate (MgATP) and the positive modulator (Mg +2) and decreases that of the inhibitor (ATP 4-).   Mg2+ or Mn2+, in contrast with Ca2+, may control not only the basal activity, but also the hormonal responsiveness of AC. A more comprehensive description of metabolic shifts requires the examination of hormonal action and metabolic feedback not only on AC, but also on insulin receptor tyrosine kinase (IRTK).  The metal dependence of IRTK, as well as the effect of changes on chelated species of ATP on this dependence, were investigated to elucidate the integrative parameters which allow coordinated hormonal, ionic and energy-charge control of the metabolic shift from glycolysis to gluconeogenesis.

 

I. Control mechanisms for secretory processes:

a) Renin release in kidney tissue.

b) HCl versus bicarbonate in gastric tissue

 


MAJOR RESEARCH INTERESTS

Metabolic Regulation: in yeast Energy transduction: in chloroplast and mitochondrial membranes, resolution and reconstitution, ATAase

Control mechanisms of enzyme-membrane systems: hormone receptors-adenylate cyclase systems in fat cells and brain membranes, insulin receptor tyrosinekinases in liver.

 

Modeling of Energy trans­duction mechanism: Hemoglobin

Areas of Specialization - Cellular actions of hormones Enzyme kinetics Cellular structure and function Membrane biochemistry of organelles, chloroplasts and mitochondria