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These interrela- tions, which are clearly of vital impor- tance In cancer therapy, are being stud- ied by several of the groups of the Re- search Program.

In view of the complexity of the topics, the work carried out in this Research Program demands the combined use of microscopic, cell-biological, biochemi- cal, and molecular biological methods and, thus, a close collaboration between the individual divisions and also with other programs within the Center.

Furthermore, affiliated with this Research Program Is a highly sophisti- cated spectroscopy department which employs the most modern analytical methods.

An approach that holds great potential is chemoprevention. The idea of this Is that suitable substances can inhibit the occurrence of cancer or cause existing tumors to stop growing or even shrink before they reach the stage of malignancy.

A prerequisite for this, however, is a knowledge of the molecular processes that one wants to manipulate. A drug that is frequently discussed in relation to chemopreven- tion is aspirin and also its relatives.

These substances are found to inhibit the development of intestinal cancer In humans. They selectively influence the communication between cells by inhibit- ing the formation of certain signal mole- cules, namely the previously mentioned prostaglandins.

With the emphasis more on treatment than prevention, cell poisons are currently being tried out. Almost all of these have serious side effects which, in combination with the problem of resistance, means that their applicability is quite limited.

But there are good prospects of better controlling the side effects, again by selectively in- fluencing the Intercellular signal trans- mission.

Hence there is reason to hope that existing cancer treatments can be- come more successful. Coordinator of the Research Program: Prof.

Friedrich Marks Divisions and their heads: Pathochemistry: Prof. Volker Kinzel Biochemistry of the Cell: Prof. Dieter Werner Biochemistry of Tissue-Specific Regulation: Prof.

Friedrich Marks Differentiation and Carcinogenesis in vitro: Prof. Norbert Fusenig Tumor Biochemistry: Prof. Dietrich Keppler Molecular Biology of Mitosis: Prof.

Herwig Ponstingl Signal Transduction and Growth Control: Dr. Peter Angel 45 3. We have developed a cell culture test which evaluates skin-irritating chemicals by measuring a central trigger mechanism in skin inflammations.

Specifically, we focus on the release of the primary me- diators of inflammation, interleukin-1 a and arachidonic acid from human kerat- inocytes.

This reaction represents a general cellular response to the effect of substances that irritate the skin. The validity of the test was confirmed in ex- periments on humans.

Clinical symp- toms of inflammation such as erythema and a disturbance in the barrier charac- teristics of the skin, as well as the local release of arachidonic acid and its re- sulting products, correlate well with the data measured in the cell culture.

On the basis of these findings, it ap- pears that this procedure may be a suit- able alternative method for certain ani- mal experiments.

It is possible to fur- ther develop this test to detect certain aspects of carcinogenic effects. From Animal Experiment to Cell Culture Test Thousands of new chemicals are devel- oped every year in the pharmaceutical, cosmetic, food, and chemical indus- tries.

Before such substances can be used, they must be tested for harmful effects, such as whether or not they cause Inflammations, have caustic ef- fects, are associated with deformities, or are carcinogenic.

The legislature re- quires various animal experiments for this very purpose. Although we cannot yet do without such experiments in light of the current state of scientific knowl- edge, their value is being examined in an increasingly critical manner.

Not only do strong public feelings with respect to animal protection and animal rights play a role here, but also economic and sci- entific arguments are being openly dis- cussed.

Testing certain chemicals for their skin-irritating properties can serve as an example. For this purpose, the procedure developed by John Draize in is still being used to this very day.

In this method test substances are placed on the skin or on the eye of a rabbit; the symptoms of an acute irrita- tion are observed over the course of several days and are classified accord- ing to their degree of severity.

Such symptoms include the reddening of the skin erythema , which can be traced to an increased blood flow in the superfi- cial vessels and swelling edema that occurs when the blood vessel walls be- come permeable and blood plasma passes into the tissue.

Although these symptoms are evaluated according to a standardized key and the test satisfies the requirements of governmental au- thorities throughout the world, doubts exist about its reliability.

For example, the reproducibility of results obtained in different laboratories and their transfer- ability to humans has been questioned.

Furthermore, the measured inflamma- tory reactions can prove to be very bur- densome for the experimental animal since they are usually accompanied by pain.

Therefore, the Draize Test stands under fire both from the public and from animal protection organizations. In par- ticular, many people no longer under- stand the need to test products for per- sonal hygiene In animal experiments.

The European Union has yielded to this public pressure In its sixth amendment to the EU Cosmetic Guidelines. As of January 1, , animal experiments are to be generally prohibited for testing 46 Research without Animal Experiments the ingredients of cosmetic substances.

However, there is reservation; scientif- ically valid alternate methods must be available. The legislature will only rec- ognize such alternate methods if they are equivalent to the "gold standard" as defined by the Draize Test.

The development of alternate methods is complicated by the fact that the in- flammatory process is a complex reac- tion in the human body.

Many types of cells work together and its course also depends on the nature of the irritant. Such a process, in all its details, cannot be mimicked in cell and tissue culture.

We be- lieve that we have identified such fun- damental processes specifically for the inflammatory reaction of the skin. On this basis, we have developed a cell culture test for chemicals that irritate the skin; it should at least partially re- place conventional animal experiments.

Since inflammatory processes In the skin and in other tissues can promote, although not cause, cancer such a test should also be able to detect certain aspects of carcinogenic effects.

In fact, the idea for the test arose out of many years of research that focused on the development of skin cancer. The Response of the Skin to Irritants As the protective organ of the body, the skin responds very sensitively to harm- ful influences In the environment.

An ef- fective barrier is found in the horny layer at the surface of the skin; It Is formed from dead epidermal cells or keratinocytes and is sealed with fat-like substances.

In the next step, white blood cells leukocytes and defense cells of the immune system are attract- ed. They all communicate by means of hormone-like signal molecules that are called mediators of inflammation.

Thereby, the characteristic symptoms of inflammation result: erythema, edema, a feeling of heat, pain, and pos- sibly pyogenesis.

Additionally, the epi- dermis is strengthened through acceler- ated cell production and cornificatlon hyperplasia.

The keratinocytes in the epidermis appear to play a key role in triggering this cascade of reactions. As a kind of interface between the body and the environment, these cells are specialized to respond to stimuli by re- leasing mediators of Inflammation and, thereby, to set in motion the protective and defense functions.

Several of these signal substances are released within a few seconds, others with some delay. For example, members of the eicosanoid family Include prosta- glandins, leukotrienes and the hydroxy- eicosatetraenoic acids.

Common to all primary mediators of inflammation is their storage within the cell so that they can be immediately released as need- ed.

It is also typical that they appear ei- ther alone or together with other active substances in all phases of the Inflam- matory process.

Their role is one of stimulating keratinocytes and other types of cells to synthesize and release additional mediators so that the cas- cade of protective and defensive reac- tions is set in motion.

The Release of Inflammation Mediators in Cell Cultures is a Measure of the Skin-Irritating Effect The release of interleukin-la and arachidonic acid from keratinocytes is a process that plays a central role in the inflammatory reaction of the skin.

Fur- thermore, it can easily be measured in tissue culture. Is it possible to assess the skin-irritating effect of chemicals in this manner?

In order to answer this question, one first had to prove that treating cell cultures with skin-irritating substances led to the release of the named mediators.

In order to avoid problems associated with the differ- ences between humans and animals from the very beginning, we chose human keratinocytes HPKII cells for these experiments.

These cells in fact respond to 15 structurally different test substances by releasing Interleukin-la, arachidonic acid and eicosanoids.

Arachidonic acid and eicosanoids were determined by using radioactively marked substances. Before the test, we grew cells in the presence of 14C arachidonic acid which was taken up 47 and incorporated into the lipids com- prising the cell membranes.

After treat- ing the cells with the test chemicals, the arachidonic acid and the eicosanoids released into the nutritive solution was extracted using organic solvents, sub- sequently separated by thin layer chromatography, and finally quantified with respect to Its radioactivity.

At the same time, the release of interleukin-1 a was measured with an Immunological enzymatic test; an enzyme coupled to a specific antibody catalyzed a color re- action that was a measure for the quan- tity of interleukin.

A vitality test was used to evaluate a cell-damaging ef- fect. The test was based on the princi- ple that only living cells can convert the colorless substance MTT to an insolu- ble, purple-black dye formazan.

The quantity of the dye corresponds to the proportion of living cells. The release of interleukin-la and arachidonic acid depends on the type and quantity of the test substance.

The greater the irritating effect and the dose of a chemical, the greater was the quantity of inflammation mediators re- leased by the cells.

Clear differences were observed in the time course of the reactions: depending on the substance, the release of mediators either oc- curred quickly, delayed or late.

A quick arachidonic acid response was not al- ways coupled with a quick interleukin- 1a response. Those substances that produced a quick cell response were not always the most effective Irritants.

Therefore, it was important to measure the release of mediators over a longer time period in order to be certain of de- tecting the irritating effect of a chemical.

In testing on animals, the irritating effect of the chemicals increases from top to bottom. Those substances marked by a star were also tested on humans in order to evaluate the reliability of the results obtained with the new method is elicited.

In this context, we noticed that the cells reacted more sensitively to the release of arachidonic acid than to Interleukin-la.

This Is probably due to the fact that the cells must be quite significantly damaged before they re- lease interleukin-la while the arachi- donic acid response requires signifi- cantly less irritation.

Irrespective of this, both quantities taken together provided the most reliable information. Confirming the Method in Humans Are the results obtained with the cell culture an adequate measure of the ef- fect of chemicals on the intact skin?

This was so even without considering the problem of data transferability from ani- mals to humans. After receiving permis- sion from an ethics commission and from governmental authorities and also in collaboration with the pharmacologi- cal division of Schering AG, we con- ducted a study to evaluate our results in humans.

For this purpose, the specific substances were applied in various concentrations to the inner side of the lower arm and the treated skin areas were covered with special chambers for an entire day.

For the next three days, a physician evaluated the visible symptoms of skin irritation, especially erythema and edema. Additionally, the extent of skin damage was measured.

An especially sensitive measure for this Is the perme- ability of the epidermis for tissue fluid, so-called transepidermal water loss. It was detected with a method that is de- scribed as the noninvasive measure- ment of evaporation among dermatolo- gists.

During a second round of treat- ment, the release of arachidonic acid, eicosanoids, and Interleukin-la in the 48 Research without Animal Experiments Fig.

Various test fields are applied to the underarm of a test subject A. In order to detect damage to the skin barrier, the outflow of water from the cutis is measured B.

In the blister, which has been artificially created by suction on the cutis, messenger sub- stances of inflammation collect as they are released by the skin cells C.

Sodium iauryl sulfate, which is contained in cleaning agents, causes a reddening of the skin D skin were quantitatively analyzed. Ran- domly selected participants in the ex- periment had the test substances ad- ministered to their skin in quantities that corresponded to the previously deter- mined individual doses for a given irri- tant.

By applying negative pressure, skin blisters were produced above the test areas. The tissue fluid that collect- ed in the blisters over time contained the mediators of inflammation that had been produced by the skin cells.

This pro- cess quantitatively determined an en- tire spectrum of eicosanoids PGDg, PGE 2 , PGFgOC, G-koto-PGFgOc, LTB 4 , 5-, 12, HETE in addition to the arachi- donic acid.

It was discovered that a skin-irritating effect caused by test chemicals always correlated with a clearly elevated eicosanoid level in the blister fluid.

Barrier disturbances of the skin which are recognizable by in- creased water permeability were only caused by strong irritants. It Is especial- ly important that the irritating effect on the intact skin predominantly agreed with the data measured in the cell cul- ture.

This was demonstrated by a com- parison of eight chemicals. However, the concentrations for Interleukin-la fluctuated so significantly from test per- son to test person that a statistically certain result could not be obtained.

Conclusion We understand our studies to be a step towards developing a cell culture test to evaluate the Inflammatory effect of chemical substances.

In this context, we proceed from a more exact knowl- edge of the fundamental molecular mechanisms Instead of measuring more or less unspecific symptoms as is done in conventional procedures.

The keratinocyte cell line HPKII , which we used, can be easily kept in cell culture without any problems.

The immunological enzymatic test and marking with isotopes are both stan- dard procedures that can be conducted in any modern laboratory.

However, both the economy and the ease of use of the test would still be significantly im- proved if one could manage without ra- dioactive substances by, for example, developing an immunological test for arachidonic acid.

If one bears in mind the complexity of the inflammatory process and the ap- parently very different mechanisms of action exhibited by inflammatory chemi- cals, the relatively high significance of such a simple test may at first glance prove surprising.

Lancet , Muller-Decker, K. WIssenschaftliche Herausfor- derungen und Perspektiven. Spektrum Aka- demischer Verlag, Berlin-Heidelberg-Oxford Fig. Karin Muller-Decker Prof.

Friedrich Marks Division of Biochemistry of Tissue-Specific Regulation In collaboration with Prof. Werner Raff Dr. Andrei Kecskes Dr.

Wolfgang Seibert Ines Zimmer Department of Clinical Pharmacology, Sobering AG, Berlin Prof. Wolf-Dieter Lehmann Central Spectroscopy, Deutsches Krebsforschungszentrum Participating scientists Dr.

Gerhard Furstenberger Thomas Heinzelmann 50 Approaches to Counteract Multidrug Resistance 3. Many malignant tu- mors initially respond to treatment with cytotoxic or cytostatic agents but devel- op subsequently a resistance to sever- al, structurally diverse anticancer agents.

This phenomenon is known as multidrug resistance of tumors. The mo- lecular mechanisms underlying the mul- tidrug resistance of malignant tumors and the development of drugs counter- acting this resistance are intensively in- vestigated by scientists in research in- stitutions, clinical research laboratories, and in the pharmaceutical industry.

Different mechanisms lead to multidrug resistance of malignant tumor cells. An increasing number of molecular mech- anisms causing multidrug resistance was recognized during the past ten years.

These distinct membrane pumps function in the transport of different drugs across the plasma membrane of tumor cells into the extracellular space.

The membrane pumps encoded by the MDR1 and MRP1 genes have different amino acid sequences and a different substrate specificity with respect to the compounds transported out of tumor cells.

Additional mechanisms of drug resistance include those caused by an increased inactivation of cytotoxic drugs by binding to glutathione or the resistance of tumor cells to the signals causing programmed cell death apop- tosis.

Resistance to structurally diverse cyto- toxic agents by ATP mediated export of Fig. Vincristine is such a product.

Along with ATP and glutathione, it can be transported out of the tumor cell by the muitidrug resistance protein MRP1 drugs from tumor cells.

It has been es- tablished that the MDR1 gene encoded P-glycoproteIn, an export pump with a molecular mass of about kllodal- ton, transports structurally diverse drugs including anthracycline-based cy- tostatic agents, vinca alcaloids, or the antimitotic drug taxol paclitaxel.

These anticancer drugs are widely used in the clinical treatment of hemat- ological cancers, ovarian cancer, and mammary cancer.

Effective concentra- tions of these anticancer agents may be prevented by the action of MDR1 P- glycoprotein. It has been recognized more recently that MDR1 P-glycopro- tein transports not only exogenous cy- totoxic drugs into the extracellular space but also a number of structurally different endogenous lipophilic com- pounds.

This membrane protein is often ex- pressed in tumors which do not have increased levels of MDR1 P-glycopro- tein and which are resistant to several anticancer drugs.

This new pump has been termed multidrug resistance pro- tein MRP1. The multidrug resistance protein MRP1 functions as an ATP dependent export pump for conjugated substanc- es.

Intracellular formation of conjugates is observed when compounds poorly soluble in water are conjugated to a negatively charged substance, such as glutathione or glucuronate, yielding conjugates of better solubility.

The func- tion and substrate specificity of the mul- tidrug resistance protein MRP1 was recognized by our group in during the search for the membrane protein Fig.

Cancer drugs, carcinogenic substances, and poisons that are taken up by the cell right can be coupled to internal cellular sub- stances conjugates and transported out of the cell by ATP-dependent pumps 52 Approaches to Counteract Multidrug Resistance which mediates the ATP dependent transport of the glutathione conjugate leukotriene C4.

MRP1 functions in ATP dependent membrane transport of neg- atively charged lipophilic compounds, such as glutathione conjugates and glu- curonate conjugates of endogenous and foreign xenobiotic substances.

Leukotriene C4 is an endogenous glu- tathione conjugate, which contributes to the development of human bronchial asthma and inflammation, and which Is transported by MRP1 with high affinity.

We have shown that conjugates of sev- eral cytotoxic agents with glutathione or glucuronate are substrates for MRP1 mediated transport across the plasma membrane of human tumor cells.

An additional substrate of MRP1 is glutath- ione disulfide GSSG which is particu- larly formed under conditions of oxida- tive stress when GSSG must be trans- ported into the extracellular space.

Thus, MRP1 had been discovered and cloned as a membrane protein asso- ciated with multidrug resistance; how- ever, only the elucidation of its function and substrate specificity has led to the molecular Identification of the ATP de- pendent transport protein for glutathi- one conjugates and glucuronides.

We have termed MRP1 a conjugate export pump in view of its capacity to transport many structurally diverse conjugates.

Cloning of a new ATP dependent ex- port pump: The apical multidrug resis- tance protein MRP2. The cell surface of epithelial cells Is characterized by specialized apical membrane domains which are oriented to the lumen of ducts and cavities.

The ATP dependent transport of conjugates across apical membrane domains of epithelial cells had been known for a long time; however, the molecular iden- tity of the corresponding transport pro- tein was unknown.

The apical multidrug re- sistance protein has been termed MRP2 or canalicular multispecific or- ganic anion transporter cMOAT. The MRP2 gene-encoded conjugate export pump in humans is a glycoprotein with a molecular mass of about kilodal- ton containing amino acids.

The substrate specificities of MRP 1 and MRP2 are rather similar in spite of the differences in the amino acid sequence of both transport proteins.

The different localization of MRP1 and MRP2 In cells containing apical membrane domains indicates a difference In the functional specialization of both conjugate export pumps.

MRP2 Is predominantly found in the apical membrane of liver cells and cells of the kidney where it contrib- utes to the transport of endogenous and xenobiotic conjugates into bile and urine, respectively.

It Is of considerable interest to establish that MRP2 contrib- utes to multidrug resistance of primary liver cancer and renal cell carcinoma.

Both types of cancer are very frequent world-wide and particularly resistant to anticancer drugs. Multidrug resistance associated with drug conjugation to glutathione and ATP dependent export of conjugates from cancer cells.

It has been known for some time that multidrug resistance of malignant tumors may be associated with enhanced cellular levels of glutath- ione and increased activities of glutathi- one-conjugating enzymes i.

Several clinical studies were conducted in an attempt to overcome the resistance to cytotoxic agents by lowering of the glutathione content of tumors and to counteract thereby the glutathione conjugation of cytotoxic agents.

As a consequence of the elucidation of the function of MRP1 and MRP2 it has become apparent that an inhibition of the export of the cyto- toxic drug conjugates with glutathione from the cell should be an additional target to enhance the antitumor action of drugs and to increase their intracellu- lar concentration by Inhibition of export.

Susan Cole, Roger Deeley, and their collaborators in Canada recently showed that the cytotoxic agent vincris- tine, which is used in the therapy of leu- kemias and other types of cancer, is transported from tumor cells to the ex- tracellular space as a complex with glu- tathione which is pumped out by MRP1.

We observed that alkylating cytotoxic agents are transported out of tumor cells by MRP1 after their intracellular conjugation with glutathione.

These re- sults Indicate that tumors expressing in- creased amounts of MRP1 in their plas- ma membrane may become resistant to a variety of structurally different cyto- toxic agents if conjugation or complex formation of the drug with glutathione proceeds at a sufficient rate.

The conjugate export pumps MRP1 and MRP2 have important physiological functions, such as the biosynthetic re- lease of the endogenous mediator leu- kotriene C4 and the excretion of biliru- bin glucuronides and estradiol glucuro- nide.

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Main Becoming Heinrich Schenker : music theory and ideology. Becoming Heinrich Schenker : music theory and ideology Morgan , Robert P.

Becoming Heinrich Schenker brings a new perspective to Schenker's theoretical work, showing that ideas characteristic of his mature theory, although in many respects fundamentally different, developed logically out of his earlier ideas.

Robert Morgan provides an introduction to Schenker's mature theory and traces its development through all of his major publications, considering each in detail and with numerous music examples.

Morgan also explores the relationship between Schenker's theory and his troubled ideology, which crucially influenced the evolution of his ideas and was heavily dependent upon both the empirical and idealist strains of contemporary German philosophical thought.

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