Degrees Offered
Doctor of Philosophy

Program Admissions Requirements
In addition to the general Graduate School admission requirements, this program has additional specific requirements.

Students with a major in the sciences, who have demonstrated above-average scholastic ability, are eligible. Minimum undergraduate course work should include: biology (8 credits), inorganic chemistry (8 credits), organic chemistry (8 credits), physics (8 credits), algebra (3-4 credits), and calculus (3-4 credits).

Fields of Research
There are eight general areas of physiology in which students may conduct research. Both human and animal models are studied.

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Current projects are directed to study the overall control of the cardiovascular system with emphasis on the neurohumoral control of arterial pressure and the interactions of the renal body fluid volume system. Regulation of cardiac output and local regulations of flow in the microcirculation is also being studied in depth. (Drs. Cowley, Greene, Liang, Lombard, and O’Connor)

  Computer Simulation of Biological Systems

Computer modeling of the complex interactions of the variables involved in the overall regulation of the cardiovascular, respiratory, and renal fluid volume control system. (Drs. Cowley, and Greene)


Current research is related to the renin-angiotensin-aldosterone system, vasopressin, and prostaglandin with emphasis on the role of these systems on the control of body fluids, electrolytes, and arterial pressure regulation. (Drs. Cowley and Raff)


Current research focuses on the identification and characterization of gene(s) involved in the development of complex disorders in humans and rat models. Research includes QTL mapping, linkage and association analysis, haplotype and linkage disequilibrium analysis of single nucleotide polymorphisms, correlation of genetic polymorphisms to gene and protein expression, and development of transgenic animal models. Projects currently study genetic risk factors for cardiovascular and renal disease, obesity, diabetes, and brain disorders. (Drs. Kwitek and Michalkiewicz)

  Molecular and Cell Biology

Research areas include cell membrane channels and their modulation by neurotransmitters, humoral agents, local factors and drugs, regulation of cardiac myosin heavy chain gene expression, cardiac hypertrophy, and DNA binding proteins. (Dr. Wakatsuki)


Research areas include the importance of the kidney in the regulation of fluid and electrolyte homeostasis and blood pressure, the importance of autocrine, paracrine, and hormonal factors in the regulation of renal tubular and vascular function, and the elucidation of the factors important in the development of renal ischemic and hypertensive disease. (Drs. Liang, Cowley and Mattson)


Aspects currently under study are: the role of the carotid chemoreceptors and medullary, pontine, and cerebellar nuclei in the control of breathing. Specific emphasis is on respiratory rhythm and pattern generation and chemosensitivity. Studies are targeted to gain insight into central and obstructive sleep apnea, the Sudden Infant Death Syndrome, and Congenital Central alveola hypoventilation. (Drs. Dwinell, ForsterPan, and Hodges)


Ongoing projects relate to the role of circulatory smooth muscle in hypertension. (Drs. Greene, Harder, and Lombard)

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  Required Courses

The emphasis of our PhD program is to provide training in whole animal integrative Physiology as well as cellular and molecular Physiology. To achieve this objective, all PhD students are required to complete a two year, 4 semester, sequence of courses.

All students are required to take General Human Physiology (08202), but to fulfill the 9 credit requirement for each semester, the student and their advisor choose courses that best meet the students career needs. Course selection can be specific to Physiology (see below) or from other MCW departments or other institutions in Milwaukee. At the completion of this sequence, all students will take the written and oral comprehensive examination. For all years of graduate study, students will enroll each semester in Special Problems in Physiology (08275). They will also enroll in a Seminar Course (08300) each summer.

  Research Requirements

Matriculating students will begin by attending the first orientation session offered in early July. For the remainder of July, the students will spend one day in the laboratory of as many faculty as possible. By August 1, they need to have selected the three laboratories they would most likely complete their research. They will then spend three months in each laboratory while taking courses. The major purpose of these rotations is to facilitate the students’ deciding the primary laboratory for their PhD research. Since students are required to complete research in whole animal, cellular, and molecular physiology and because of the highly collaborative research by our faculty, most students will conduct research in the laboratories of their advisor and also in other laboratories. Students will thus have broad exposure and experience in the basic techniques of integrated physiology, molecular genetics and physiological genomics. At the end of each three-month rotation, the students summarize orally for ten minutes their experiences to the faculty.

  Overall Course Requirements

A requirement of this program is to fulfill two credits in Bioethics by completing Course (10222) Ethics and Integrity in Science and Course (10444) Research Ethics Discussion Series. For course descriptions of 10222 and 10444, see listing within Bioethics Program.


08203 Complement to General Human Physiology. 1 credit. Concurrent enrollment in 08202 General Human Physiology.
Basic functions of cells, tissues, and organ systems are discussed weekly in relation with the General Human Physiology course. Short individual oral examinations are used as a starting point for discussions that emphasize problem solving and integrative thinking.

08204 Graduate Human Physiology. 4 credits
This course provides the fundamental aspects of: 1) cell membrane transport, 2)smooth, skeletal, and cardiac contractile mechanisms and excitation-contraction coupling, 3) principles of synaptic transmission, neurotransmitters, and neuromodulators, 4) respiratory mechanics, gas exchange in the lung, and control of breathing, 5) cardiac electrophysiology, hemodynamics, nervous and humoral control of the heart and cardiac output, 6) renal tubular transport, glomerular filtration, and regulation of sodium and water balance, 7) neural and humoral control of gastrointestinal absorption and motility, and 8) endocrinology including pituitary, adrenal cortical, thyroid, pancreatic and male and female reproductive organs. Material will be covered primarily in lecture format by expert physiologists in each respective area. There will be three multiple choice examinations.

08210 Endocrine Regulation and Common Disease. 1 credit. Prerequisite 08202.
This course presents current knowledge regarding endocrine regulation. Introduction by instructor, student presentations, and discussion formats will be utilized to develop skills for extraction of information, critical thinking, and oral and written communication of primary endocrine literature. Each session consists of three parts: (a) instructor introduction; (b) presentation of a review article by a student; and (c) presentation of a research article by a student. Each student is expected to study these articles and actively participate in critical evaluation of the paper.

08212 Critical Reading in Respiratory Physiology. 1 credit.
This course includes critical reading and communication of primary literature in respiratory physiology. Discussion and student presentation formats will be utilized to develop skills for extraction of information, critical thinking, and oral and written communication of primary respirator physiology literature.

08223 Bioinformatics–Tools and Tactics. 1 credit.
This introductory course is targeted at anyone who needs to understand and use online bioinformatics tools and analyses as part of their research activities. It focuses on providing the background understanding and practical methods suitable for the functional annotation and interpretation of proteins or genes derived from microarray analysis, proteomics, genome-wide association studies and similar experimental techniques. The online resources covered will be centered around mammalian biology (human, rat, mouse) but resources from other species may be covered where appropriate. The course will be a mixture of lecture/discussion combined with practical sessions in the computer lab.

08225 Molecular Biology for Physiologists. 1 credit.
This course will provide a basic overview of molecular and cellular process that is important for the basic researcher to function in the lab. If applicable, techniques that can be used to study these processes will be described to provide relevance to the textbook material.

08229 Essential Physiological Genomics. 2 credits.
This course covers genome sequence, functional genomic analysis, genome and gene manipulation, and grant writing. The students will learn about the latest advances in the field of physiological genomics, how to apply genomic approaches to study complex physiological problems and how to develop a grant proposal.

08230 Physiological Genomes. 5 credits.
This course will cover topics in Physiological Genomics at an advanced level emphasizing the tools and techniques that are available to investigators exploring the relationship between genotype and phenotype. Material will be selected to emphasize
high throughput screening and Bioinformatics techniques. Specific examples of applications of physiological genomics to important research problems will be discussed. Students will acquire the expertise required to develop a research proposal and will participate in a mock study section to witness the process by which grants are reviewed.

08253 Advanced Renal Physiology. 1 credit. Prerequisite: 08202.
This course is an in-depth survey of selected important topics in renal physiology that will be taught at an advanced level appropriate for a senior graduate student. The material in this course will emphasize classical and modern research approaches and techniques, original research papers, and the development of oral presentation skills. The knowledge gained from this course should be sufficient for a doctoral student in physiology.

08255 Advanced Cardiovascular Physiology. 1 credit. Prerequisite: 08202.
This course will use a guided approach to reading of selected contemporary literature in cardiovascular physiology to explore the relationships between genetic and environmental influences on the development and progression of major cardiovascular diseases including Hypertension, Heart failure and Metabolic Syndrome. The course will combine didactic lectures with class discussion and will help students to develop skills for extraction of information, critical thinking, and oral and written communication on concepts in Cardiovascular Physiology.

08273 Special Topics in Neuroscience. 1 credit. Prerequisite: 01207.
The objective of this course is to critically evaluate, discuss and debate the strengths and weaknesses of a particular contemporary literature neuroscience study and understand the study’s broader relation to neuroscience and how this hypothesis changed the field. Each semester the course will choose a new overlying theme (e.g. neural plasticity, rhythm generation, sleep, learning and memory, development, locomotion, neuro-computational studies, etc.) and applicable literature.

08275 Special Problems in Physiology. Variable credits.
Readings and/or research under direction of a faculty member in a specialized field of physiology. Under specific circumstances, may be substituted for formal courses.

08284 Computational Methods in Biomedical Research. 3 credits. Prerequisite proficient understanding of undergraduate calculus,
familiarity with introductory concepts of linear algebra, and access to any conveniently available language or programming environment
(such as Fortran, C, Matlab, or Mathematica). Nearly every area of modern research in biology and biomedical science is targeted in one way or another on gaining quantitative understanding of the behavior of biological systems. This course will focus on practical techniques for simulation and analysis of biological systems, developed largely through application-driven examples. Examples will be developed to a depth at which models will be used to analyze real biological or physiological data. To accomplish this, the important details of the underlying biological systems must be described along with a complete step-by step development of model assumptions, the resulting equations, and (when necessary) computer code.

08295 Reading and Research. 1-9 credits.

08299 Master’s Thesis. 6 credits.

08301 Seminar. 1 credit.
Presentations by visiting scholars, resident investigators, and graduate students on topics of physiological research. All full-time graduate students are expected to attend.

08399 Doctoral Dissertation. 9 credits.

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MCW Physiology News

Dr. Mingyu Liang honored with 2017 Distinguished Lectureship in Physiological Genomics Research

May 26 - Dr. Mingyu Liang, MD, PhD, director of the Center of Systems Molecular Medicine and professor & eminent scholar in physiology, has been named the 2017 Distinguished Lectureship Awardee in Physiological Genomics Research. Dr. Liang gave a presentation titled, The Essential Role of Physiological Genomics in Precision Medicine, at the Experimental Biology Conference in Chicago