MODULATION OF CALCIUM ACTIVATED POTASSIUM CHANNELS T. M. WEIGER, A. HERMANN, I. B. LEVITAN

Size: px
Start display at page:

Download "MODULATION OF CALCIUM ACTIVATED POTASSIUM CHANNELS T. M. WEIGER, A. HERMANN, I. B. LEVITAN"

Transcription

1 Selected abstracts: MODULATION OF CALCIUM ACTIVATED POTASSIUM CHANNELS T. M. WEIGER, A. HERMANN, I. B. LEVITAN Potassium currents play a critical role in action potential repolarization, setting of the resting membrane potential, control of neuronal firing rates, and regulation of neurotransmitter release. The diversity of the potassium channels that generate these currents is nothing less than staggering. This diversity is generated by multiple genes (as many as 100 and perhaps more in some creatures) encoding the pore-forming channel subunits, alternative splicing of channel gene transcripts, formation of heteromultimeric channels, participation of auxiliary (non-pore-forming) and other subunits, and modulation of channel properties by posttranslational modifications and other mechanisms. Prominent among the potassium channels are several families of calcium activated potassium channels, which are highly selective for potassium ions as their charge carrier, and require intracellular calcium for channel gating. The modulation of one of these families, that of the large conductance calcium activated and voltage-dependent potassium channels, has been especially widely studied. In this review we discuss a few selected examples of the modulation of these channels, to illustrate some of the molecular mechanisms and physiological consequences of ion channel modulation. Figure Putative structures of BK channel subunits. The pore-forming subunits of the Slowpoke family of BK channels are large proteins (~1,200 amino acids) that resemble other voltage-dependent potassium channels in having six membrane-spanning domains (S1 - S6), with a pore region between S5 and S6. An additional membrane-spanning domain (S0) places the amino terminal outside the plasma membrane. Most notable is the extended carboxyl terminal tail domain, comprising about two thirds of the subunit protein sequence. It includes a negatively charged region (the so-called calcium bowl) that has been implicated in calcium binding, and is the site of interaction with several channel modulatory proteins including protein kinases. The auxiliary subunits are small proteins (~200 amino acids) with two membrane-spanning domains (T1 and T2)., potential sites for N-linked glycosylation. Modified from Vergara et al., WEIGER, T., HERMANN, A. & LEVITAN, I. B. (2002). Modulation of calcium-activated potassium channels. J. Comp. Physiol. A (invited Review), 188,

2 MOLEKÜLSONDEN ZUR ERFORSCHUNG VON IONENKANÄLEN T. M. WEIGER, T. LANGER, A. HERMANN Ionenkanäle sind wesentliche Bestandteile jeder lebenden Zelle bei Mensch, Tier und Pflanzen. Sie sind verantwortlich für bioelektrische Vorgänge in unserem Körper, wie zum Beispiel für die Entstehung der elektrischen Aktivität bei Sinnes-, Nerven-, und Muskelzellen, die Volumenregulation oder die Steuerung des Salz-/Wasserhaushalts. Ionenkanäle sind winzige Nanoporen die man in allen Zellmembranen, wie der Plasmamembran um Zellen, bei Mitochondrien, dem Endoplasmatischen Reticulum oder in der Membran des Zellkerns, findet, Zelluläre Membranen bestehen aus Doppelschichten von Lipiden (fettähnliche Stoffe) die für die meisten geladenen Teilchen (Ionen) wie Natrium, Kalium, Calcium oder Chlorid unpassierbar sind. Durch Kanäle, die in diese Lipdschicht eingelagert sind, können Ionen ausgetauscht werden. Auf den ersten Blick sind Ionenkanäle einfache, wassergefüllte Poren die Ionen, durch die Membran schleusen. Aber ungleich einfachen Löchern in der Membran sind Ionenkanäle tatsächlich sehr kompliziert gebaute Eiweißmoleküle (Proteine). Mittels elektrophysiologischer und molekulargenetischer Verfahren kann man heute die Funktion und Struktur dieser Kanalproteine erforschen. Ionenkanäle können entweder offen oder geschlossen sein. Sie sind demnach winzige molekulare Schalter die im offenen Zustand Ionen entlang ihrem Konzentrationsgradienten fließen lassen. Zudem sind viele Ionenkanäle hoch selektiv, das heißt, sie lassen nur eine bestimmte Sorte von Ionen durch. Diese Eigenschaft hängt mit der molekularen Struktur der Kanäle zusammen, wobei eine enge Stelle im Kanal, der sog. Selektivitätsfilter eine wichtige Rolle spielt. Große Bedeutung haben Wirkstoffe, die Ionenkanäle blockieren oder modulieren können. In der Medizin werden solche Substanzen zu mannigfaltigen Therapien eingesetzt, wie z. B. bei multipler Sklerose, bei Herzrhythmusstörungen, bei Epilepsie etc. An Hand von einfach gebauten Molekülen und deren Varianten, wie den natürlich vorkommenden Polyaminen und ihren Analoga, den Diaminen, wird gezeigt, wie diese Substanzen quasi als Sonden zur Erforschung von Kaliumkanälen eingesetzt werden können. PUT SPD SP 1,12DD Computersimulation der Hydrathüllen (korbartige Strukturen) die Putrescin (PUT), Spermidine (SPD), Spermin (SP) und 1,12-Diaminododecan (1,12-DD) umgeben. Beachten Sie, daß alle Moleküle mit Ausnahme von 1,12-Diaminododecan vollständig von einer Hydrathülle umschlossen sind. 1,12-Diaminododecan kann daher leichter eine seine beiden, kleinen, endständigen Hydrathüllen abstreifen und in den Ionenkanal gelangen.

3 Schematische Darstellung des Block eines Kalium-Kanals durch Tetraethylammonium und 1,12- Diaminododecan. Die Abbildung zeigt von links nach rechts einen Kaliumkanal durch den Kaliumionen (rot) ungehindert fliessen können (links), ein Tetraethylammonium Molekül (grün, Mitte) das nur ein kurzes Stück in den Kanal von außen eindringen kann, im Kanal stecken bleibt und ihn somit blockiert, und rechts 1,12-Diaminododecan das die Wasserhülle von einer seiner beiden positiv geladenen Aminogruppen abstreifen kann und in gestreckter Form tief in den Kanal eindringt. Die zweite Wasserhülle von 1,12-Diaminododecan wird wahrscheinlich nicht abgestreift, das Molekül ragt auf Grund seiner Länge bis in den Vorhof des Kanals. (Graphik: Peter Steinbacher). WEIGER, T., LANGER, T., & HERMANN, A. (2002). Molekülsonden zur Erforschung von Ionenkanälen. Biologie i. u. Zeit, 2, COMPLEMENTARY DISTRIBUTION OF NADPH DIAPHORASE AND L ARGININE IN THE SNAIL NERVOUS SYSTEM M. XIE, A. HERMANN, H. H. KERSCHBAUM Since the interneuronal messenger nitric oxide (NO) can not be stored in neurons, the regulation of the NO producing enzyme, nitric oxide synthase (NOS), is crucial. Neuronal NOS metabolizes L - arginine to nitric oxide (NO) and L citrulline in a Ca 2+ dependent manner. Therefore, availability of L arginine to NOS may modulate NO production. Using NADPH diaphorase histochemistry to visualize putative NO producing cells and immunocytochemistry, we showed that the distribution of L arginine immunoreactive neurons correlates well with those of NADPH positive neurons in cerebral ganglia of the pulmonate, Helix pomatia. However, substrate and enzyme were visualized in separate but adjacent neurons. Following elevation of intracellular Ca 2+ by the Ca 2+ ionophore, ionomycin, or by a high K + solution, the number of L - citrulline immunoreactive neurons in mesocerebrum and pedal lobe increased up to ten fold. Preincubation of ganglia with the NOS inhibitor, N G -nitro-l-arginine, prevented ionomycin or high K + solution induced L - citrulline synthesis. Most L - citrulline immunoreactive neurons revealed NADPH diaphorase staining. In conclusion, these

4 experiments indicate a complementary distribution of NOS and L arginine and suggests an unknown signaling pathway between neurons to maintain L arginine and NO homeostasis. XIE, M., HERMANN, A. & KERSCHBAUM, H. H. (2002). Complementary distribution of NADPH-diaphorase and L-arginine in the snail nervous system. Cell Tissue Res., 307, NITRIC OXIDE AND CGMP MEDIATED MODULATION OF CA AND K CA - CONDUCTANCES IN SNAIL NEURONS S. SCHROFNER, A. ZSOMBOK, A. HERMANN, H. H. KERSCHBAUM The mechanism of nitric oxide (NO) - dependent modulation of Ca 2+ - and Ca 2+ - activated potassium (K Ca ) conductance was studied in identified subesophageal neurons of the pulmonate snail, Helix pomatia. K Ca is voltage as well as Ca 2+ - dependent and sensitive to charybdotoxin (CTX), Ba 2+, and tetraethylammonium (TEA), but insensitive to aminopyridine (4 AP). Thus, the K Ca current shows similarities to a large-conductance K Ca channel (BK(Ca)). NO - donors (sodium nitroprusside, SNP; S nitro N acetylpenicillamine, SNAP) slowly decreased the BK current amplitude. Decline of the current amplitude by NO - donors was qualitatively mimicked by a membrane permeable cgmp analogue (dibutyryl cgmp, db- cgmp). Methylene blue, an inhibitor of guanylyl cyclase, or erytho 9 (2 hydroxyl 3 nonyl) adenine (EHNA), an inhibitor of the cgmp stimulated phosphodiesterase 2, inhibited NO - donor induced decrease of BK - current. Amplitude of L type Ca 2+ - current either did not respond to NO donors or increased following application of NO donors. Exposure of neurons to db cgmp increased the current amplitude. In EHNA pretreated neurons, amplitude of L type Ca 2+ - current did not respond to NO or cgmp. The experiments demonstrate that the NO mediated affects on Ca 2+ - and K Ca conductance in snail neurons dependent on generation of cgmp and subsequent cgmp SCHROFNER, S., ZSOMBOK, A., HERMANN, A. & KERSCHBAUM, H.H. (2004). Nitric oxide decreases a calcium activated potassium current via activation of phosphodiesterase 2 in Helix U cells. Brain Res., 999, MODULATION OF CALCIUM CHANNELS PHOSPHORYLATION BY NITRIC OXIDE IN SNAIL NEURONS A. ZSOMBOK, S. SCHROFNER, A. HERMANN, H. H. KERSCHBAUM The different responses of cells to nitric oxide (NO) exposure in many cases depends on the direct interaction of NO with metal or thiol containing proteins, like enzymes or ion channels, which enhances or attenuates the activity of these proteins. Although NO modulates L type calcium channel activity in vertebrate tissue, it appears not to directly affect the L type calcium current in snail neurons. This difference could result from an insensitivity of calcium channels to NO or NO dependent processes or may be caused by a simultaneous phosphorylation and dephosphorylation of calcium channels. We recently reported NO donors to increase the intracellular concentration of cgmp in Helix ganglia (Huang et al., Brain Res. 1998, 780:329) and to increase excitability by depressing a calcium-activated potassium current (Zsombok et al., 2000, Neurosci Lett, 295: 85).

5 In the present study, we investigated the effect of the membrane permeant 3,5 cyclic guanosine monophosphate (cgmp) analogue, dibutyryl cgmp (dbcgmp) and the NO donor sodium nitroprusside (SNP) on the L type voltage-activated Ca 2+ current in identified neurons in the subesophageal of the terrestrial snail, Helix pomatia. We observed that application of db cgmp resulted in a transient increase of the calcium current by about 30%, whereas in most neurons SNP had no effect on the current amplitude. Treatment of ganglia with the kinase inhibitor, H-7, abolished the enhancing effect of dbcgmp. Furthermore, pretreatment of ganglia with H-7 decreased the calcium current following application of dbcgmp and the NO donor SNP by about 25%. The phosphatase inhibitors okadaic acid and calyculin increased the Ca 2+ current following application of dbcgmp and SNP by about 30%. Our experiments indicate that NO modulates phosphorylation of calcium channels via activation of a cgmp dependent transduction pathway. ZSOMBOK, A., SCHROFNER, S., HERMANN, A & KERSCHBAUM, H.H. (2005). A cgmpdependent cascade enhances an L-type-like Ca 2+ current in identified snail neurons. Brain Res., 1032, LOCALISATION OF ANNEXINS IN THE RETINA OF RAINBOW TROUT: LIGHT- AND ELECTRON MICROSCOPICAL INVESTIGATIONS ZAUNREITER M., BRANDSTÄTTER R., DONATO, R., A. HERMANN We present a first description of annexin immunoreactivity within neural cells of the teleost retina. Antibodies against annexins V and VI were used in light and electron miscoscopic sections of light and dark adapted retinae. Strong immunoreactivity could be found in retinal layers with high synaptic input, such as the outer and inner plexiform layers and dendritic regions within the inner plexiform layer; in cells that are involved in negative feedback control, such as horizontal and amacrine cells; in the membrane metabolism of photoreceptor outer segments; and in close relation to cytoskeletal components. Our findings suggest that both, annexins V and VI are involved in the regulation of transmitter release, especially of transmitters that are not directly involved in phototransduction, and with structures that support the morphological changes involved in light and dark adaptation. ZAUNREITER, M., BRANDSTÄTTER, R., DONATO, R. & HERMANN, A. (2005). Localisation of annexins in the retina of the rainbow trout - light- and electron microscopical investigations. Brain Res., 1032, ANTIPROLIFERATIVE PROPERTIES OF PADMA LAX AND ITS COMPONENTS GINGER AND ELECAMPANE SEBASTIAN HOFBAUERA, VERENA KAINZ, LEOPOLD GOLSER, MICHAELA KLAPPACHER, TOBIAS KIESSLICH, WILHELM HEIDEGGER, BARBARA KRAMMER, ANTON HERMANN & THOMAS M. WEIGER Padma Lax (PL) is a multi-component herbal laxative, derived from traditional Tibetan medicine. It has been used in the treatment of constipation dominant irritable bowel syndrome. Beyond its purgative and bowel-regulating properties we found it to exhibit antiproliferative properties. C6 tumor cells were incubated with either an ethanolic or aqueous extract of PL. Cell proliferation, cell cycle, percentage of apoptotic cells, caspase-3/-7 activity as well as

6 mitochondrial membrane potential were determined. Ethanolic extracts of PL inhibited cell proliferation in a dose- and time-dependent manner (half max concentration: μg/ml after 48 h of incubation). Aqueous extracts were less effective. Ginger and elecampane were the active components of PL in respect to its antiproliferative action and were found to act synergistically. Supplementing the culture medium with polyamines could not override the cytostatic action of PL. Incubation of C6 cells with PL in the presence of catalase proved that the PL effect was specific and not due to oxidative stress. PL had no effects on the cell cycle at a low dose but arrested cells in G1 at high concentrations. Reduction of cell numbers was found to be due to apoptosis. The caspase-3/-7 pathway was not involved in the PL-induced cell death. However, mitochondrial membrane potential was lost during the course of incubation with PL indicating a mitochondrial- but not caspase-mediated induction of apoptosis. PL exhibits antiproliferative properties which may be beneficial to prevent constipation-related cancer. This study may also contribute to a future development of a new herbal-based antiproliferative treatment. HOFBAUER, S., KAINZ, V., GOLSER, L., KLAPPACHER, M., KIESSLICH, T., HEIDEGGER, W., KRAMMER, B., HERMANN, A. & WEIGER, T.M. (2006). Antiproliferative properties of Padma Lax and ist components ginger and elecampane. Forschende Komplementärmedizin, 13, HYPOTONICITY AND ETHANOL MODULATE BK CHANNEL ACTIVITY AND CHLORIDE CURRENTS IN GH4/C1 PITUITARY TUMOUR CELLS M. JAKAB, S. SCHMIDT, M. GRUNDBICHLER, M. PAULMICHL, A. HERMANN, T. WEIGER & M. RITTER Description of the effects of hypotonic cell swelling and ethanol onmaxi Ca2+-activated K+ channel (BK channel) activity and Cl) channel activity in GH4/C1 pituitary tumour cells. Whole cell-, cell attached- and outside-out patch clamp measurements, fluorescence (fluo-3) measurements of intracellular Ca2+ concentration, cell size video monitoring. GH4/C1 pituitary tumour cells respond to both hypotonicity and ethanol with cell swelling which is followed by a regulatory volume decrease (RVD). Tetraethylammonium and 4,4 diisothiocyanatostilbene-2,2 -disulphonic acid (DIDS) induced cell swelling per se and inhibited hypotonicity induced RVD. Ethanol-induced swelling is paralleled by an increase in the intracellular Ca2+ concentration and augmented by DIDS. BK channel activation by hypotonicity and ethanol is demonstrated in patch clamp experiments both in intact cells (cell attached configuration) and a subset of excised membrane patches (outside-out configuration). Cell swelling and addition of ionomycin under isotonic conditions leads to the activation of outwardly rectifying Cl) currents with time dependent activation at positive potentials. In GH4/C1 cells both hypotonicity and ethanol lead to cell swelling, RVD and to activation of BK channels. The hypotonicity-induced BK channel activation can also be observed in cell free outside-out patches. Hypotonicity, but not ethanol leads to the activation of Cl) channels. JAKAB, M., SCHMIDT S., GRUNDBICHLER, M., PAULMICHL, M., HERMANN, A., WEIGER, T. & RITTER, M. (2006). Hypertonicity and ethanol modulate BK channel activity and chloride currents in GH4/CI pitiutary tumor cells. Acta Physiol. 187,

7 POTASSIUM CHANNEL BLOCKERS QUINIDINE AND CAESIUM HALT CELL PROLIFERATION IN C6 GLIOMA CELLSVIA A POLYAMINE-DEPENDENT MECHANISM T.M. WEIGER, S. COLOMBATTO, V. KAINZ, W. HEIDEGGER, M.A. GRILLO & A. HERMANN Potassium (K + ) channels are ubiquitous to cells and serve essential functions in physiology and pathophysiology. K + channel blockers have been shown to block tumour growth by arresting cells at the G0/G1 checkpoint of the cell cycle. We investigated the effect of quinidine and caesium (Cs+) on cell proliferation, LDH (lactate dehydrogenase) release, free internal calcium (Ca 2+ ), membrane potential, polyamine concentration, ODC (ornithine decarboxylase) activity and polyamine uptake in C6 glioma cells. The EC50 for reducing cell proliferation was 112 μm for quinidine, whereas Cs + was less effective with an EC50 of 4.75 mm. LDH release was augmented by quinidine. It caused a transient increase in free internal Ca 2+ but decreased Ca 2+ after a 48 h incubation period. Further 300 μm quinidine depolarized the cell membrane in a similar range as did 30 mm KCl. Quinidine decreased cellular putrescine beyond detection levels while spermidine and spermine remained unaffected. ODC activity was reduced. Addition of putrescine could not override the antiproliferative effect owing to a reduced activity of the polyamine transporter. Our study indicates that the antiproliferative effect of quinidine is not due to a simple membrane depolarization but is caused by a block of ODC activity. WEIGER, T.M., COLOMBATTO, S., KAINZ, V., HEIDEGGER, W., GRILLO, M.A. & HERMANN, A. (2007). Potassium channel blockers quinidine and caesium halt cell proliferation in C6 glioma cells via a polyamine-dependent mechanism. Biochemical Society Transactions, 35-2, MODULATION OF POTASSIUM CHANNELS BY POLYAMINES THOMAS M. WEIGER AND ANTON HERMANN Polyamines modulate a large variety of potassium channels. Not only Kir channels which have received most attention are affected bypolyamines but also calcium activated potassium channels like BK channels, the delayed rectifier, KCNQ channels, and TASK channels are modulated by polyamines. In all cases polyamines block potassium channels. The order of potency is spermine > spermidine> putrescine. In most of these channels polyamines act from the intracellular side of the channels and show no blocking properties when applied from the extracellular side except TASK-3 channels. The polyamine action on potassium channels has important bearings on the discharge activity or secretion in excitable cells. WEIGER, T. & HERMANN, A. (2009). Modulation of potassium channels by polyamines, In: Biological Aspects of Biogenic Amines, Polyamines and Conjugates, Ed. G. Dandrifosse, Transworld Research Network, ETHANOL AND POLYAMINES IN BRAIN THOMAS M. WEIGER AND ANTON HERMANN In this review we discuss the role of the polyamines putrescine, spermidine and spermine which are ubiquitous constituents in all eukaryotic cells, during acute and chronic ethanol exposure as well as after ethanol withdrawal in brain. Ethanol as well as polyamines modulate the same set

8 p(open) of ion channels with the NMDA receptor being a main target. Ethanol exposure or withdrawal is responsible for changes in polyamine content via modulation of ornithine decarboxylase, the rate limiting enzyme for polyamine synthesis. Alterations of polyamine concentrations in turn affect ion channels interacting with polyamines in addition to the direct effect of ethanol on these ion channels. Any increase or decrease in ion channel activity impacts electrical signalling in the brain and subsequently behaviour. A better understanding of these mechanisms concerning ethanol, polyamines and ion channel activity may generate improved treatment options for alcoholics. Not only ethanol but also acetaldehyde, the first by-product of ethanol metabolism, may be involved in the modulation of polyamines as well as ion channels, a field that is largely ignored by recent research. WEIGER, T.M. & HERMANN, A. (2009). Ethanol and polyamines in brain. In: Biologically Active Amines and Related Enzymes: Biochemical, Physiological and Clinical Aspects, Ed. A. Toninello, Transworld Research Network, HYDROGEN SULFIDE INCREASES CALCIUM-ACTIVATED POTASSIUM (BK) CHANNEL ACTIVITY OF RAT PITUITARY TUMOR CELLS GUZEL F. SITDIKOVA, THOMAS M. WEIGER & ANTON HERMANN Hydrogen sulfide (H2S) is the third gasotransmitter found to be produced endogenously in living cells to exert physiological functions. Large conductance (maxi) calcium-activated potassium channels (BK) which play an important role in the regulation of electrical activity in many cells are targets of gasotransmitters. We examined the modulating action of H2S on BK channels from rat GH3 pituitary tumor cells using patch clamp techniques. Application of sodium hydrogen sulfide as H2S donor to the bath solution in whole cell experiments caused an increase of calcium-activated potassium outward currents. In single channel recordings H2S increased BK channel activity in a concentration-dependent manner. H2S induced a reversible increase in channel open probability in a voltage-dependent, but calcium independent manner. The reducing agent dithiothreitol prevented the increase of open probability by H2S whereas the oxidizing agent thimerosal increased channel open probability in the presence of H2S. Our data show that H2S augments BK channel activity and this effect can be linked to its reducing action on sulfhydryl groups of the channel protein. SITDIKOVA, G.F., WEIGER, T.M. & HERMANN, A. (2010). Hydrogen sulphide increases calcium-activated potassium (BK) channel activity of rat pituitary tumor cells. Pflügers Arch - Eur J Physiol 459, ms control NaHS 300 µm wash 10 pa sec

9 GASOTRANSMITTER - GASE ALS ZELLULÄRE SIGNALMOLEKÜLE ANTON HERMANN, GUZEL F. SITDIKOVA, UND THOMAS M. WEIGER Gase, wie Stickoxid (NO), Kohlenmonoxid (CO) oder Schwefelwasserstoff (H 2 S) sind als umweltbelastende Industrieabfallstoffe und als sehr giftige Substanzen bekannt. Dass Gase mit physiologischer Wirksamkeit in unseren Körper vorkommen sollten, war zunächst schwer vorstellbar. Nun wurde aber gefunden, dass diese Gase in allen Organismen von Bakterien bis zum Menschen vorkommen, in deren Zellen produziert werden und wichtige Funktionen erfüllen. HERMANN, A., SITDIKOVA, G.F. & WEIGER, T.M. (2010).Gase als zelluläre Signalstoffe. Biologie i.u. Zeit, 3, GIFTIGE GEISTER VON ANTON HERMANN, GUZEL F. SITDIKOVA UND THOMAS M. WEIGER Stickstoffmonoxid, Kohlenmonoxid und Schwefelwasserstoff gelten als hochtoxisch. Umso erstaunlicher ist, dass der menschliche Körper diese Gase selbst produziert: Sie dienen im Organismus als universelle Botenmoleküle. Die Biologen Anton Hermann, Guzel F. Sitdikova und Thomas M. Weiger erforschen die vielfältigen Wirkungen der Gasotransmitter etwa beim Lernen oder bei neurodegenerativen Erkrankungen. Ein»animalischer Geist«durchströmt nach Ansicht der alten Griechen unsere Adern. AlsLebenskraft wirke dieses Pneuma vor allem im Gehirn, im Herzen und in der Leber. Heute wissen wir es besser und lehnen solche Vorstellungen als esoterisch ab. Vielleicht etwas vorschnell denn geisterhaft wirkende Substanzen konnten Wissenschaftler in den letzten Jahrzehnten tatsächlich aufspüren: Die Gase Stickstoffmonoxid, Kohlenmonoxid und Schwefelwasserstoff kontrollieren wichtige Stoffwechselprozesse in Zellen und Organen unseres Körpers. HERMANN, A., SITDIKOVA, G.F. & WEIGER, T.M. (2011). Giftige Geister. Gehirn & Geist 5, 28-35, Spektrum Verlag.

10 S100 CALCIUM-BINDING PROTEINS AND ION CHANNELS ANTON HERMANN 1*, ROSARIO DONATO 2, THOMAS M. WEIGER 1 AND WALTER J. CHAZIN 3 S100 Ca 2+ -binding proteins have been associated with a multitude of intracellular Ca 2+ - mediated functions including regulation of the cell cycle, cell differentiation, cell motility and apoptosis, modulation of membrane-cytoskeletal interactions, transduction of intracellular Ca 2+ signals, and in mediating learning and memory. S100 proteins are fine tuned to read the intracellular free Ca 2+ concentration and affect protein phosphorylation, which makes them candidates to modulate certain ion channels and neuronal electrical behaviour. Certain S100s are secreted from cells and are found in extracellular fluids where they exert unique extracellular functions. In addition to their neurotrophic activity, certain S100 proteins modulate neuronal electrical discharge activity and appear to act directly on ion channels. The first reports regarding these effects suggested S100-mediated alterations in Ca 2+ fluxes, K + currents and neuronal discharge activity. Recent reports revealed direct and indirect interactions with Ca 2+, K +, Cl - and ligand activated channels. This review focuses on studies of the physical and functional interactions of S100 proteins and ion channels. HERMANN, A., DONATO, R., WEIGER, T.M. &.CHAZIN, W.J. (2012). S100 Calcium Binding Proteins and Ion Channels. Front. Pharmacol. 3 (67), p. 1-10, doi: /fpharm , Review. Online:http://www.frontiersin.org/Journal/Abstract.aspx?s=762&name=neuropharmacology& ART_DOI= /fphar BK CHANNELS - FOCUS ON POLYAMINES, ETHANOL/ACETALDEHYDE AND HYDROGEN SULFIDE (H2S) HERMANN ANTON, SITDIKOVA GUZEL F. & WEIGER THOMAS M. Calcium-activated potassium channels are expressed in a great variety of tissues from bacteria to men. The channels constitute a link between cellular activity, calcium signalling and metabolism. The importance of the channels results from their unique adaptability, modulating capacity and versatile physiology. To study these channels many scientists have been attracted to this field who created a great wealth of detailed insight into structure and function of these channels. In this chapter we will describe techniques to record maxi calcium-activated potassium (BK) channels, delineate properties and functions of BK channels and we will focus on some aspects of channel modulation by polyamines, ethanol /acetaldehyde and by the gasotransmitter - hydrogen sulfide. HERMANN, A., SITDIKOVA, G.F. & WEIGER, T.M. (2012). BK Channels Focus on Polyamines, Ethanol/Acetaldehyde and Hydrogen Sulfide (H2S), In Patch Clamp Technique, InTech, Ed. F. S. Kaneez, p , ISBN: , Review. Online:

11 GASOTRANSMITTERS: PHYSIOLOGY AND PATHOPHYSIOLOGY ANTON HERMANN, GUZEL F. SITDIKOVA, THOMAS M. WEIGER - EDITORS Since the epochal discovery of the radical and highly toxic gas nitric oxide (NO) as signaling molecule two other not less toxic gases carbon monoxide (CO) and hydrogen sulfide (H 2 S) - have been found to be also involved in a plethora of physiological and pathophysiological functions. The gases termed Gasotransmitters play an increasingly important role in understanding how signalling into cells and between cells is modulated and fine tuned The advent of gasotransmitters has profoundly changed our way of thinking about biosynthesis, liberation, storage and action mechanisms by cellular signaling. In recent years an impressive amount of new data distributed in the literature has been generated. For this book we have asked distinguished colleagues in the field to summarize and review important biological, pharmacological and medical functions and their implications as well as methods for the detection of gasotransmitters. MODULATED BY GASOTRANSMITTERS - BK CHANNELS HERMANN ANTON, SITDIKOVA GUZEL F. & WEIGER THOMAS M. Calcium-activated potassium BK channels interconnect cellular activity, calcium signaling and cell metabolism. Major virtues of these channels are their adaptability to different functions, their versatile physiology and their capacity being modulated. The channels are present in a large variety of cells and organs in different forms of life from bacteria to men. Scientists attracted to these channels have produced a great wealth of information regarding their structure and function. Mutations at channels proteins are involved in a number of diseases (channelopathies), like diabetes, epilepsy or heart failure. The gasotransmitters NO, CO and H 2 S all act on BK channels directly or indirectly via signaling pathways. In this chapter we will briefly summarize some of the basic properties of BK channels and focus on aspects of BK channel modulation by gasotransmitters and their implications in physiology and pathophyiology. HERMANN, A., SITDIKOVA, G.F. & WEIGER, T.M. (2012). Gasotransmitters: Physiology and Pathophysiology (book), Eds. A. Hermann. G. Sitdikova & T. Weiger, Springer Press, Heidelberg, Germany, ISBN: e-book citation: HERMANN, A., SITDIKOVA, G.F. & WEIGER, T.M. (2012). Modulated by Gasotransmitters BK channels, In Gasotransmitters: Physiology and Pathophysiology, Eds. A. Hermann et al., Springer Press, Heidelberg, Germany, p , Review.

12 CELL PROLIFERATION, POTASSIUM CHANNELS, POLYAMINES AND THEIR INTERACTIONS WEIGER THOMAS M. & HERMANN A. Polyamines, which are obligatory molecules involved in cell cycling and proliferation are subject to a change in their free intracellular concentrations during the cell cycle. Potassium (K + ) channels are also considered, but less well recognized, to be necessary for cell proliferation by either hyperpolarizing or depolarizing cells during the cell cycle. A block of polyamine synthesis as well as block or knockout of K + channels can halt cell proliferation. K + channels like BK (maxi calcium (Ca 2+ ) activated K + ), Kir (inward rectifier), M-type K + - and TASK (two-pore domain K + ) channels or the delayed rectifier K + channels are modulated in their electricalproperties by polyamines. Polyamines are most effective in blocking these channels when applied from the intracellular faces of these channels except for TAKS channels where they act from the extracellular side. Quinidine a general K + channel blocker was found to reduce putrescine concentrations, to block the ornithine decarboxylase and to halt cell proliferation. From these results the question arises if there is an interaction between polyamines, K + channels and proliferation. It might be speculated that a decrease of intracellular polyamines allows more K + channels to be active thus inducing hyperpolarization while an increase of the polyamine concentration may block K + channel activity leading to depolarization of the membrane potential. On the other hand, a block or a deletion of K + channels may cause a decrease of the polyamine concentration in cells. More research is needed to test these hypotheses. WEIGER, T.M. & HERMANN, A. (2013). Cell proliferation, potassium channels, polyamines and their interactions. Amino Acids. DOI /s Link:http://www.springer.com/home?SGWID= &aqId= &download=1&checkval=cf85b2afe38a972b9efeb3232c1cefb1 SCREENING OF ANALGESIC AND ANTI-INFLAMMATORY ACTIVITIES FOR TWO LIBYAN MEDICINAL PLANTS: HELIANTHEMUM LIPPII AND LAUNAEA RESIDIFOLIA NOURI B. ERMELI, SAMI G. ALSABRI, SALAH M. BENSABER, SALAH B. MOHAMED, ABDULMOTTALEB A. ZETRINI, KHALED M. ABURAS, SAFA R. FITOURI, MOUSA I. JAEDA, IBRAHIM A. MREMA, A. HERMANN & ABDUL M. GBAJ1 Natural products are often a source for bioactive compounds which have great potential for developing novel therapeutic agents. In this study, two Libyan medicinal plants Helianthemum lippii (H. lippii) and Launaea residifolia (L. residifolia) were collected from El-Jabel El- Garbi (Gharian) in the Spring season (2010). They were extracted successively by using microwave technique with three different solvents of different polarities. The analgesic activity of these plant extracts was evaluated using the hot-plate method and the anti-inflammatory activity was evaluated using Carrageenen-induced paw edema method. The methanol and chloroform extracts exhibited significant analgesic activity at the doses tested while the petroleum ether extracts of both plants did not show any significant effect. In addition, the anti-inflammatory activities of various extracts showed a significant percentage inhibition of paw edema for H. lippii extracts in methanol and chloroform but not in petroleum ether. Moreover, the results exhibit different percentage inhibitions of paw edema for L. residifolia extracts in methanol, chloroform and petroleum ether. The analgesic and anti-inflammatory effects produced by the

13 extracts may be attributed individually or collectively to the flavonoids and tannins. H. lippii and L. residifolia can be introduced as new plant sources for analgesics and anti inflammatory agents. The methanolic and chloroform extracts of both plants showed a significant analgesic activity due to an increase in the reaction time (p<0.05) in comparison to controls, where codeine was used as standard analgesic drug. The petroleum ether extracts did not show any activity as analgesic. The anti-inflammatory activity was also evaluated using the Carrageenaninduced paw edema method, the methanolic extract of H. lippii (53.42%) exhibit activity comparable to that of aspirin standard antiinflammatory drug (62.28%) with no significant difference, while the petroleum ether extract of L. residifolia (31.65%) exhibited a moderate anti-inflammatory activity (p<0.01) in comparison with aspirin as a standard. The chloroform extract of H. lippii, the methanol and chloroform extracts of L. residifolia exhibited a weak inhibitory effect on paw edema volume with percentage inhibition of (23.60%, p<0.05), (16.45%, p>0.05) and (14.56%, p>0.05) respectively compared to control. ERMELI, N.B., ALSABRI, S.G., BENSABER, S.M., MOHAMED, S.B., ZETRINI, A.A., ABURAS, K.M., FITOURI, S.R., JAEDA, M.I., MREMA, I.A., HERMANN, A.& GBAJ, A.M. (2012). Screening of analgesic and anti-inflammatory activities for two Libyan medicinal plants: Helianthemum lippii and Launaea residifolia. Journal of Chemical and Pharmaceutical Research, 4(9): BRCA1 MUTATION DETECTION USING FLUORESCENT HYBRIDIZATION PROBES AND MELTING CURVES SAFA R. FITOURI, NOURI B. ERMELI, SALAH M. BENSABER, MOUSA I. JAEDA, IBRAHIM A. MREMA, ANTON HERMANN & ABDUL M. GBAJ The BRCA1 (Breast Cancer Anti-estrogen resistance-1), early-onset gene is expressed in cells of breast and other tissue and helps to repair damaged DNA or destroy cells in cases DNA cannot be repaired. When the BRCA1 gene is damaged, then the DNA is not repaired appropriately and this enhances the risk for cancer. Fluorescence and UV-visible thermal studies were performed for WT (wild type) and MT (mutant type targets) full systems. The target DNAs used were in the form of short oligonucleotides, genomic DNA. The probe system was used for detection of WT and SNP alleles of human BRCA1 [( , G T) and ( , G T)]. The Cy5 dye attached to a probe oligonucleotide (10-mer) undergoes a fluorescence intensity change on hybridisation of the probe to the WT compared to MT targets. Our results indicate that the system consisting of the target sequence and the one probe oligonucleotides bearing the Cy5 dye assemble correctly at the specified target. Once the full system (probe and target) is arranged under suitable conditions, a red-shift emission and change in fluorescence intensity are seen at a suitable wavelength. Thermal studies also showed significant differences in Tm between WT and MT. The results suggest that the differences in the fluorescence intensity at 665 nm and the spectrophotometric Tm(s) for the WT and MT can be attributed to the type of binding of the probe to the target. The systems were sensitive to single nucleotide polymorphisms and this may help in high throughput applications in genetic testing and molecular diagnostics. FITOURI, S.R., ERMELI, N.B., BENSABER, S.M., JAEDA M.I., MREMA, I.A., HERMANN, A. & GBAJ, A.M. (2013). BRCA1 Mutation Detection Using Fluorescent Hybridization Probes and Melting Curves. Journal of Life Sciences, USA, 7-3, ISSN

14 CHEMICAL SYNTHESIS, MOLECULAR MODELLING AND EVALUATION OF ANTICANCER ACTIVITY OF SOME PYRAZOLIDONE SCHIFF'S BASE DERIVATIVES S. M. BENSABER, H. A. ALLAFE, Z. S. ABOOD,, N. B. ERMELI, S. B. MOHAMED, A. EL- ZITRINI, S. G. ALSABRI, I. A. MREMA, M. ERHUMA, ANTON HERMANN, M. I. JOAED & A. M. GBAJ A series of novel 3-pyrazolinone Schiff`s base derivatives (MCS 02-13) were designed and synthesized by microwave chemical synthesis. Their purity was confirmed by melting point and HPLC and their chemical structures were determined by FT-IR, UV, 1H and 13C-NMR spectroscopic techniques. In silico the synthesized compounds have been docked to in the thymidine phosphorylase active site using molecular modeling programs. The compounds were tested in vitro on calf thymus DNA to study the interaction with DNA using a Cary 5000 UV\VIS\NIR spectrophotometer. Some of these compounds showed close match interaction with DNA. The compounds were also studied on angiogenic enzyme thymidine DNA phosphorylase (TP, E.C ), carcinoma cell lines including both human breast (MCF-7) and human lung cell lines (A549). The lead compound in the series, MCS 2, exhibited inhibition of thymidine phosphorylase in the micro molar range (IC50 of 28 ± 2 µm) and was able to retard growing of breast carcinoma cells. Our results indicate that 3-pyrazolinone Schiff`s base derivatives are promising lead compounds for the development of more active antitumor agents and exhibit their highest cytotoxic effect on breast carcinoma cell line. BENSABER, S.M., ALLAFE, H.A., ABOOD, Z.S., ERMELI, N.B., MOHAMED, S.B., EL- ZITRINI, A., ALSABRI, S.G., MREMA, I.A., ERHUMA, M., HERMANN, A., JOAED, M.I. & GBAJ, A.M. (2013). Chemical synthesis, molecular modelling and evaluation of anticancer activity of some pyrazolidone Schiff's base derivatives. (in press).

PART I: Neurons and the Nerve Impulse

PART I: Neurons and the Nerve Impulse PART I: Neurons and the Nerve Impulse Identify each of the labeled structures of the neuron below. A. B. C. D. E. F. G. Identify each of the labeled structures of the neuron below. A. dendrites B. nucleus

More information

Chapter 8. Summary and Perspectives

Chapter 8. Summary and Perspectives Chapter 8 Summary and Perspectives 131 Chapter 8 Summary Overexpression of the multidrug resistance protein MRP1 confer multidrug resistance (MDR) to cancer cells. The contents of this thesis describe

More information

Mechanism of short-term ERK activation by electromagnetic fields at mobile phone frequencies. Biochemistry Journal. August 1, 2007 405, pp.

Mechanism of short-term ERK activation by electromagnetic fields at mobile phone frequencies. Biochemistry Journal. August 1, 2007 405, pp. Mechanism of short-term ERK activation by electromagnetic fields at mobile phone frequencies 1 Biochemistry Journal August 1, 2007 405, pp. 559 568 Joseph Friedman, Sarah Kraus, Yirmi Hauptman, Yoni Schiff

More information

INTRODUCTION TO HORMONES

INTRODUCTION TO HORMONES INTRODUCTION TO HORMONES UNIVERSITY OF PNG SCHOOL OF MEDICINE AND HEALTH SCIENCES DISCIPLINE OF BIOCHEMISTRY & MOLECULAR BIOLOGY PBL MBBS II SEMINAR VJ Temple What are hormones? Cells in multi-cellular

More information

The diagram below summarizes the effects of the compounds that cells use to regulate their own metabolism.

The diagram below summarizes the effects of the compounds that cells use to regulate their own metabolism. Regulation of carbohydrate metabolism Intracellular metabolic regulators Each of the control point steps in the carbohydrate metabolic pathways in effect regulates itself by responding to molecules that

More information

Chapter-21b: Hormones and Receptors

Chapter-21b: Hormones and Receptors 1 hapter-21b: Hormones and Receptors Hormone classes Hormones are classified according to the distance over which they act. 1. Autocrine hormones --- act on the same cell that released them. Interleukin-2

More information

Protein Function. After the Folding. Lecture 3

Protein Function. After the Folding. Lecture 3 Protein Function After the Folding Lecture 3 Gene to gene product (protein) Protein folding of nascent polypeptide chain - Immediate folding amplification Proteins mediate virtually all cellular functions

More information

Student name ID # 2. (4 pts) What is the terminal electron acceptor in respiration? In photosynthesis? O2, NADP+

Student name ID # 2. (4 pts) What is the terminal electron acceptor in respiration? In photosynthesis? O2, NADP+ 1. Membrane transport. A. (4 pts) What ion couples primary and secondary active transport in animal cells? What ion serves the same function in plant cells? Na+, H+ 2. (4 pts) What is the terminal electron

More information

Resting membrane potential ~ -70mV - Membrane is polarized

Resting membrane potential ~ -70mV - Membrane is polarized Resting membrane potential ~ -70mV - Membrane is polarized (ie) Electrical charge on the outside of the membrane is positive while the electrical charge on the inside of the membrane is negative Changes

More information

NO CALCULATORS OR CELL PHONES ALLOWED

NO CALCULATORS OR CELL PHONES ALLOWED Biol 205 Exam 1 TEST FORM A Spring 2008 NAME Fill out both sides of the Scantron Sheet. On Side 2 be sure to indicate that you have TEST FORM A The answers to Part I should be placed on the SCANTRON SHEET.

More information

ANIMATED NEUROSCIENCE

ANIMATED NEUROSCIENCE ANIMATED NEUROSCIENCE and the Action of Nicotine, Cocaine, and Marijuana in the Brain Te a c h e r s G u i d e Films for the Humanities & Sciences Background Information This program, made entirely of

More information

Uses of Flow Cytometry

Uses of Flow Cytometry Uses of Flow Cytometry 1. Multicolour analysis... 2 2. Cell Cycle and Proliferation... 3 a. Analysis of Cellular DNA Content... 4 b. Cell Proliferation Assays... 5 3. Immunology... 6 4. Apoptosis... 7

More information

Unit 2 Metabolism and Survival Summary

Unit 2 Metabolism and Survival Summary Unit 2 Metabolism and Survival Summary 1 Metabolism pathways and their control (a) Introduction to metabolic pathways This involves integrated and controlled pathways of enzymecatalysed reactions within

More information

Cell Biology Questions and Learning Objectives

Cell Biology Questions and Learning Objectives Cell Biology Questions and Learning Objectives (with hypothetical learning materials that might populate the objective) The topics and central questions listed here are typical for an introductory undergraduate

More information

Hormones & Chemical Signaling

Hormones & Chemical Signaling Hormones & Chemical Signaling Part 2 modulation of signal pathways and hormone classification & function How are these pathways controlled? Receptors are proteins! Subject to Specificity of binding Competition

More information

D. Vitamin D. 1. Two main forms; vitamin D2 and D3

D. Vitamin D. 1. Two main forms; vitamin D2 and D3 D. Vitamin D. Two main forms; vitamin D2 and D3 H H D3 - Cholecalciferol D2 - Ergocalciferol Technically, vitamin D is not a vitamin. It is the name given to a group of fat-soluble prohormones (substances

More information

2006 7.012 Problem Set 6 KEY

2006 7.012 Problem Set 6 KEY 2006 7.012 Problem Set 6 KEY ** Due before 5 PM on WEDNESDAY, November 22, 2006. ** Turn answers in to the box outside of 68-120. PLEASE WRITE YOUR ANSWERS ON THIS PRINTOUT. 1. You create an artificial

More information

* Is chemical energy potential or kinetic energy? The position of what is storing energy?

* Is chemical energy potential or kinetic energy? The position of what is storing energy? Biology 1406 Exam 2 - Metabolism Chs. 5, 6 and 7 energy - capacity to do work 5.10 kinetic energy - energy of motion : light, electrical, thermal, mechanical potential energy - energy of position or stored

More information

PHC 313 The 7 th. Lecture. Adrenergic Agents

PHC 313 The 7 th. Lecture. Adrenergic Agents PHC 313 The 7 th. Lecture Adrenergic Agents Introduction Introduction Adrenergic agents are a broad class of agents employed in the treatment of many disorders. They are those chemical agents that exert

More information

SICKLE CELL ANEMIA & THE HEMOGLOBIN GENE TEACHER S GUIDE

SICKLE CELL ANEMIA & THE HEMOGLOBIN GENE TEACHER S GUIDE AP Biology Date SICKLE CELL ANEMIA & THE HEMOGLOBIN GENE TEACHER S GUIDE LEARNING OBJECTIVES Students will gain an appreciation of the physical effects of sickle cell anemia, its prevalence in the population,

More information

Bi 360: Midterm Review

Bi 360: Midterm Review Bi 360: Midterm Review Basic Neurobiology 1) Many axons are surrounded by a fatty insulating sheath called myelin, which is interrupted at regular intervals at the Nodes of Ranvier, where the action potential

More information

Alcohol Use Dates Back 7,000 to 10,000 Years. Though Scientists Still Debate the Mechanisms of Hangovers. Proposed Causes of Hangovers

Alcohol Use Dates Back 7,000 to 10,000 Years. Though Scientists Still Debate the Mechanisms of Hangovers. Proposed Causes of Hangovers Alcohol Use Dates Back 7,000 to 10,000 Years Though Scientists Still Debate the Mechanisms of Hangovers Proposed Causes of Hangovers Acute ethanol withdrawal Ethanol can alleviate symptoms Acetaldehyde

More information

BIOLOGICAL MEMBRANES: FUNCTIONS, STRUCTURES & TRANSPORT

BIOLOGICAL MEMBRANES: FUNCTIONS, STRUCTURES & TRANSPORT BIOLOGICAL MEMBRANES: FUNCTIONS, STRUCTURES & TRANSPORT UNIVERSITY OF PNG SCHOOL OF MEDICINE AND HEALTH SCIENCES DISCIPLINE OF BIOCHEMISTRY AND MOLECULAR BIOLOGY BMLS II / B Pharm II / BDS II VJ Temple

More information

Mechanisms of Hormonal Action Bryant Miles

Mechanisms of Hormonal Action Bryant Miles Mechanisms of ormonal Action Bryant Miles Multicellular organisms need to coordinate metabolic activities. Complex signaling systems have evolved using chemicals called hormones to regulate cellular activities.

More information

Twincore - Zentrum für Experimentelle und Klinische Infektionsforschung Institut für Molekulare Bakteriologie

Twincore - Zentrum für Experimentelle und Klinische Infektionsforschung Institut für Molekulare Bakteriologie Twincore - Zentrum für Experimentelle und Klinische Infektionsforschung Institut für Molekulare Bakteriologie 0 HELMHOLTZ I ZENTRUM FÜR INFEKTIONSFORSCHUNG Technische Universität Braunschweig Institut

More information

Actions of Hormones on Target Cells Page 1. Actions of Hormones on Target Cells Page 2. Goals/ What You Need to Know Goals What You Need to Know

Actions of Hormones on Target Cells Page 1. Actions of Hormones on Target Cells Page 2. Goals/ What You Need to Know Goals What You Need to Know Actions of Hormones on Target Cells Graphics are used with permission of: Pearson Education Inc., publishing as Benjamin Cummings (http://www.aw-bc.com) Page 1. Actions of Hormones on Target Cells Hormones

More information

Diabetes and Insulin Signaling

Diabetes and Insulin Signaling Diabetes and Insulin Signaling NATIONAL CENTER FOR CASE STUDY TEACHING IN SCIENCE by Kristy J. Wilson School of Mathematics and Sciences Marian University, Indianapolis, IN Part I Research Orientation

More information

2007 7.013 Problem Set 1 KEY

2007 7.013 Problem Set 1 KEY 2007 7.013 Problem Set 1 KEY Due before 5 PM on FRIDAY, February 16, 2007. Turn answers in to the box outside of 68-120. PLEASE WRITE YOUR ANSWERS ON THIS PRINTOUT. 1. Where in a eukaryotic cell do you

More information

Lecture 8. Protein Trafficking/Targeting. Protein targeting is necessary for proteins that are destined to work outside the cytoplasm.

Lecture 8. Protein Trafficking/Targeting. Protein targeting is necessary for proteins that are destined to work outside the cytoplasm. Protein Trafficking/Targeting (8.1) Lecture 8 Protein Trafficking/Targeting Protein targeting is necessary for proteins that are destined to work outside the cytoplasm. Protein targeting is more complex

More information

Common Course Topics Biology 1406: Cell and Molecular Biology

Common Course Topics Biology 1406: Cell and Molecular Biology Common Course Topics Biology 1406: Cell and Molecular Biology 1. Introduction to biology --the scientific study of organisms --properties of life --assumptions, methods and limitations of science --underlying

More information

Slide 1: Introduction Introduce the purpose of your presentation. Indicate that you will explain how the brain basically works and how and where

Slide 1: Introduction Introduce the purpose of your presentation. Indicate that you will explain how the brain basically works and how and where Slide 1: Introduction Introduce the purpose of your presentation. Indicate that you will explain how the brain basically works and how and where drugs such as heroin and cocaine work in the brain. Tell

More information

treatments) worked by killing cancerous cells using chemo or radiotherapy. While these techniques can

treatments) worked by killing cancerous cells using chemo or radiotherapy. While these techniques can Shristi Pandey Genomics and Medicine Winter 2011 Prof. Doug Brutlag Chronic Myeloid Leukemia: A look into how genomics is changing the way we treat Cancer. Until the late 1990s, nearly all treatment methods

More information

Anatomy and Physiology Placement Exam 2 Practice with Answers at End!

Anatomy and Physiology Placement Exam 2 Practice with Answers at End! Anatomy and Physiology Placement Exam 2 Practice with Answers at End! General Chemical Principles 1. bonds are characterized by the sharing of electrons between the participating atoms. a. hydrogen b.

More information

October 23, 2014. How EMR causes illnesses and what to do about it. prof. em. Martin Pall Washington State University

October 23, 2014. How EMR causes illnesses and what to do about it. prof. em. Martin Pall Washington State University October 23, 2014 How EMR causes illnesses and what to do about it prof. em. Martin Pall Washington State University 1 2 Problem 1: How can electromagnetic fields (EMFs) impact our biology and medicine

More information

Tibor G. Szántó Medical and Health Science Center, University of Debrecen Department of Biophysics and Cell Biology

Tibor G. Szántó Medical and Health Science Center, University of Debrecen Department of Biophysics and Cell Biology Resting potential, action potential and electrical excitibility. Measurement of membrane potential. Tibor G. Szántó Medical and Health Science Center, University of Debrecen Department of Biophysics and

More information

CELL MEMBRANES, TRANSPORT, and COMMUNICATION. Teacher Packet

CELL MEMBRANES, TRANSPORT, and COMMUNICATION. Teacher Packet AP * BIOLOGY CELL MEMBRANES, TRANSPORT, and COMMUNICATION Teacher Packet AP* is a trademark of the College Entrance Examination Board. The College Entrance Examination Board was not involved in the production

More information

Genetic research identifies novel pathway leading to myocardial

Genetic research identifies novel pathway leading to myocardial Press Release Embargo: 10 November 2013 at 1800 London time / 1300 US Eastern Time Genetic research identifies novel pathway leading to myocardial infarction Starting with a severely affected family, a

More information

BCOR 11 Exploring Biology Exam # 2

BCOR 11 Exploring Biology Exam # 2 BCOR 11 Exploring Biology Exam # 2 Name Section For this Multiple Choice Exam you should record your choice of the best answer for each question on the SCANTRON sheet. You must use a number 2 pencil for

More information

AP BIOLOGY 2008 SCORING GUIDELINES

AP BIOLOGY 2008 SCORING GUIDELINES AP BIOLOGY 2008 SCORING GUIDELINES Question 1 1. The physical structure of a protein often reflects and affects its function. (a) Describe THREE types of chemical bonds/interactions found in proteins.

More information

Chapter 9 Mitochondrial Structure and Function

Chapter 9 Mitochondrial Structure and Function Chapter 9 Mitochondrial Structure and Function 1 2 3 Structure and function Oxidative phosphorylation and ATP Synthesis Peroxisome Overview 2 Mitochondria have characteristic morphologies despite variable

More information

Parts of the Nerve Cell and Their Functions

Parts of the Nerve Cell and Their Functions Parts of the Nerve Cell and Their Functions Silvia Helena Cardoso, PhD [ 1. Cell body] [2. Neuronal membrane] [3. Dendrites] [4. Axon] [5. Nerve ending] 1. Cell body The cell body (soma) is the factory

More information

Biological Membranes. Impermeable lipid bilayer membrane. Protein Channels and Pores

Biological Membranes. Impermeable lipid bilayer membrane. Protein Channels and Pores Biological Membranes Impermeable lipid bilayer membrane Protein Channels and Pores 1 Biological Membranes Are Barriers for Ions and Large Polar Molecules The Cell. A Molecular Approach. G.M. Cooper, R.E.

More information

The Need for a PARP in vivo Pharmacodynamic Assay

The Need for a PARP in vivo Pharmacodynamic Assay The Need for a PARP in vivo Pharmacodynamic Assay Jay George, Ph.D., Chief Scientific Officer, Trevigen, Inc., Gaithersburg, MD For further infomation, please contact: William Booth, Ph.D. Tel: +44 (0)1235

More information

Translation Study Guide

Translation Study Guide Translation Study Guide This study guide is a written version of the material you have seen presented in the replication unit. In translation, the cell uses the genetic information contained in mrna to

More information

Notch 1 -dependent regulation of cell fate in colorectal cancer

Notch 1 -dependent regulation of cell fate in colorectal cancer Notch 1 -dependent regulation of cell fate in colorectal cancer Referees: PD Dr. Tobias Dick Prof. Dr. Wilfried Roth http://d-nb.info/1057851272 CONTENTS Summary 1 Zusammenfassung 2 1 INTRODUCTION 3 1.1

More information

Neurophysiology. 2.1 Equilibrium Potential

Neurophysiology. 2.1 Equilibrium Potential 2 Neurophysiology 2.1 Equilibrium Potential An understanding of the concepts of electrical and chemical forces that act on ions, electrochemical equilibrium, and equilibrium potential is a powerful tool

More information

Genetics Lecture Notes 7.03 2005. Lectures 1 2

Genetics Lecture Notes 7.03 2005. Lectures 1 2 Genetics Lecture Notes 7.03 2005 Lectures 1 2 Lecture 1 We will begin this course with the question: What is a gene? This question will take us four lectures to answer because there are actually several

More information

PRESTWICK ACADEMY NATIONAL 5 BIOLOGY CELL BIOLOGY SUMMARY

PRESTWICK ACADEMY NATIONAL 5 BIOLOGY CELL BIOLOGY SUMMARY Name PRESTWICK ACADEMY NATIONAL 5 BIOLOGY CELL BIOLOGY SUMMARY Cell Structure Identify animal, plant, fungal and bacterial cell ultrastructure and know the structures functions. Plant cell Animal cell

More information

Search Engines Chapter 2 Architecture. 14.4.2011 Felix Naumann

Search Engines Chapter 2 Architecture. 14.4.2011 Felix Naumann Search Engines Chapter 2 Architecture 14.4.2011 Felix Naumann Overview 2 Basic Building Blocks Indexing Text Acquisition Text Transformation Index Creation Querying User Interaction Ranking Evaluation

More information

Common Course Topics Biology 1414: Introduction to Biotechnology I

Common Course Topics Biology 1414: Introduction to Biotechnology I Common Course Topics Biology 1414: Introduction to Biotechnology I Assumptions Students may be enrolled in this course for several reasons; they are enrolled in the Biotechnology Program, they need a science

More information

Diabetes and Drug Development

Diabetes and Drug Development Diabetes and Drug Development Metabolic Disfunction Leads to Multiple Diseases Hypertension ( blood pressure) Metabolic Syndrome (Syndrome X) LDL HDL Lipoproteins Triglycerides FFA Hyperinsulinemia Insulin

More information

Structure and Function of DNA

Structure and Function of DNA Structure and Function of DNA DNA and RNA Structure DNA and RNA are nucleic acids. They consist of chemical units called nucleotides. The nucleotides are joined by a sugar-phosphate backbone. The four

More information

Ions cannot cross membranes. Ions move through pores

Ions cannot cross membranes. Ions move through pores Ions cannot cross membranes Membranes are lipid bilayers Nonpolar tails Polar head Fig 3-1 Because of the charged nature of ions, they cannot cross a lipid bilayer. The ion and its cloud of polarized water

More information

Mit einem Auge auf den mathema/schen Horizont: Was der Lehrer braucht für die Zukun= seiner Schüler

Mit einem Auge auf den mathema/schen Horizont: Was der Lehrer braucht für die Zukun= seiner Schüler Mit einem Auge auf den mathema/schen Horizont: Was der Lehrer braucht für die Zukun= seiner Schüler Deborah Löwenberg Ball und Hyman Bass University of Michigan U.S.A. 43. Jahrestagung für DidakEk der

More information

Copyright 2000-2003 Mark Brandt, Ph.D. 54

Copyright 2000-2003 Mark Brandt, Ph.D. 54 Pyruvate Oxidation Overview of pyruvate metabolism Pyruvate can be produced in a variety of ways. It is an end product of glycolysis, and can be derived from lactate taken up from the environment (or,

More information

THE EUKARYOTIC CELL CYCLE AND CANCER: IN DEPTH

THE EUKARYOTIC CELL CYCLE AND CANCER: IN DEPTH THE EUKARYOTIC CELL CYCLE AND CANCER: IN DEPTH ABOUT THIS WORKSHEET This worksheet complements the Click and Learn The Eukaryotic Cell Cycle and Cancer and is intended as an in-depth examination of the

More information

Sickle cell anemia: Altered beta chain Single AA change (#6 Glu to Val) Consequence: Protein polymerizes Change in RBC shape ---> phenotypes

Sickle cell anemia: Altered beta chain Single AA change (#6 Glu to Val) Consequence: Protein polymerizes Change in RBC shape ---> phenotypes Protein Structure Polypeptide: Protein: Therefore: Example: Single chain of amino acids 1 or more polypeptide chains All polypeptides are proteins Some proteins contain >1 polypeptide Hemoglobin (O 2 binding

More information

Version 1 2015. Module guide. Preliminary document. International Master Program Cardiovascular Science University of Göttingen

Version 1 2015. Module guide. Preliminary document. International Master Program Cardiovascular Science University of Göttingen Version 1 2015 Module guide International Master Program Cardiovascular Science University of Göttingen Part 1 Theoretical modules Synopsis The Master program Cardiovascular Science contains four theoretical

More information

Keystone Review Practice Test Module A Cells and Cell Processes. 1. Which characteristic is shared by all prokaryotes and eukaryotes?

Keystone Review Practice Test Module A Cells and Cell Processes. 1. Which characteristic is shared by all prokaryotes and eukaryotes? Keystone Review Practice Test Module A Cells and Cell Processes 1. Which characteristic is shared by all prokaryotes and eukaryotes? a. Ability to store hereditary information b. Use of organelles to control

More information

Lesson 3 Reading Material: Oncogenes and Tumor Suppressor Genes

Lesson 3 Reading Material: Oncogenes and Tumor Suppressor Genes Lesson 3 Reading Material: Oncogenes and Tumor Suppressor Genes Becoming a cancer cell isn t easy One of the fundamental molecular characteristics of cancer is that it does not develop all at once, but

More information

Regulation of enzyme activity

Regulation of enzyme activity 1 Regulation of enzyme activity Regulation of enzyme activity is important to coordinate the different metabolic processes. It is also important for homeostasis i.e. to maintain the internal environment

More information

Mechanisms of action of AEDs

Mechanisms of action of AEDs Mechanisms of action of AEDs Wolfgang Löscher Department of Pharmacology, Toxicology and Pharmacy University of Veterinary Medicine Hannover, Germany and Center for Systems Neuroscience Hannover, Germany

More information

Consultation on Drugs for Rare Diseases

Consultation on Drugs for Rare Diseases Consultation on Drugs for Rare Diseases Rare Disease Day Symposium Hannover Medical School (MHH), Solidarity Rare but Strong Together Roland Seifert, MHH @ Dear Sir, I am writing hoping that there might

More information

STANDARD 2 Students will demonstrate appropriate safety procedures and equipment use in the laboratory.

STANDARD 2 Students will demonstrate appropriate safety procedures and equipment use in the laboratory. BIOTECHNOLOGY Levels: 11-12 Units of Credit: 1.0 CIP Code: 51.1201 Prerequisite: Biology or Chemistry Skill Certificates: #708 COURSE DESCRIPTION is an exploratory course designed to create an awareness

More information

1.1.2. thebiotutor. AS Biology OCR. Unit F211: Cells, Exchange & Transport. Module 1.2 Cell Membranes. Notes & Questions.

1.1.2. thebiotutor. AS Biology OCR. Unit F211: Cells, Exchange & Transport. Module 1.2 Cell Membranes. Notes & Questions. thebiotutor AS Biology OCR Unit F211: Cells, Exchange & Transport Module 1.2 Cell Membranes Notes & Questions Andy Todd 1 Outline the roles of membranes within cells and at the surface of cells. The main

More information

Application Note No. 2 / July 2012. Quantitative Assessment of Cell Quality, Viability and Proliferation. System

Application Note No. 2 / July 2012. Quantitative Assessment of Cell Quality, Viability and Proliferation. System Application Note No. 2 / July 2012 Quantitative Assessment of Cell Quality, Viability and Proliferation System Quantitative Assessment of Cell Quality, Viability and Proliferation Introduction In vitro

More information

Module 3 Questions. 7. Chemotaxis is an example of signal transduction. Explain, with the use of diagrams.

Module 3 Questions. 7. Chemotaxis is an example of signal transduction. Explain, with the use of diagrams. Module 3 Questions Section 1. Essay and Short Answers. Use diagrams wherever possible 1. With the use of a diagram, provide an overview of the general regulation strategies available to a bacterial cell.

More information

Lecture 1 MODULE 3 GENE EXPRESSION AND REGULATION OF GENE EXPRESSION. Professor Bharat Patel Office: Science 2, 2.36 Email: b.patel@griffith.edu.

Lecture 1 MODULE 3 GENE EXPRESSION AND REGULATION OF GENE EXPRESSION. Professor Bharat Patel Office: Science 2, 2.36 Email: b.patel@griffith.edu. Lecture 1 MODULE 3 GENE EXPRESSION AND REGULATION OF GENE EXPRESSION Professor Bharat Patel Office: Science 2, 2.36 Email: b.patel@griffith.edu.au What is Gene Expression & Gene Regulation? 1. Gene Expression

More information

Energy Metabolism and Mitochondria

Energy Metabolism and Mitochondria Energy Metabolism and Mitochondria Date: September 2, 2005 * Time: 9:40 am- 10:30 am * Room: G-202 Biomolecular Building Lecturer: Mohanish Deshmukh 7109E Neuroscience Research Building mohanish@med.unc.edu

More information

Endocrine Responses to Resistance Exercise

Endocrine Responses to Resistance Exercise chapter 3 Endocrine Responses to Resistance Exercise Chapter Objectives Understand basic concepts of endocrinology. Explain the physiological roles of anabolic hormones. Describe hormonal responses to

More information

Name: Teacher: Olsen Hour:

Name: Teacher: Olsen Hour: Name: Teacher: Olsen Hour: The Nervous System: Part 1 Textbook p216-225 41 In all exercises, quizzes and tests in this class, always answer in your own words. That is the only way that you can show that

More information

ATOMS AND BONDS. Bonds

ATOMS AND BONDS. Bonds ATOMS AND BONDS Atoms of elements are the simplest units of organization in the natural world. Atoms consist of protons (positive charge), neutrons (neutral charge) and electrons (negative charge). The

More information

7 Answers to end-of-chapter questions

7 Answers to end-of-chapter questions 7 Answers to end-of-chapter questions Multiple choice questions 1 B 2 B 3 A 4 B 5 A 6 D 7 C 8 C 9 B 10 B Structured questions 11 a i Maintenance of a constant internal environment within set limits i Concentration

More information

博 士 論 文 ( 要 約 ) A study on enzymatic synthesis of. stable cyclized peptides which. inhibit protein-protein interactions

博 士 論 文 ( 要 約 ) A study on enzymatic synthesis of. stable cyclized peptides which. inhibit protein-protein interactions 博 士 論 文 ( 要 約 ) 論 文 題 目 A study on enzymatic synthesis of stable cyclized peptides which inhibit protein-protein interactions ( 蛋 白 質 間 相 互 作 用 を 阻 害 する 安 定 な 環 状 化 ペプチドの 酵 素 合 成 に 関 する 研 究 ) 氏 名 張 静 1

More information

CelaSYS Customized PEGylation Solutions

CelaSYS Customized PEGylation Solutions CelaSYS Customized PEGylation Solutions Neue Ära der Wirkstoffoptimierung - New Era of Drug Optimization Die celares GmbH verfügt über exzellentes fachliches Know-how in den Bereichen Synthesechemie, Biotechnologie

More information

Date: Student Name: Teacher Name: Jared George. Score: 1) A cell with 1% solute concentration is placed in a beaker with a 5% solute concentration.

Date: Student Name: Teacher Name: Jared George. Score: 1) A cell with 1% solute concentration is placed in a beaker with a 5% solute concentration. Biology Keystone (PA Core) Quiz Homeostasis and Transport - (BIO.A.4.1.1 ) Plasma Membrane, (BIO.A.4.1.2 ) Transport Mechanisms, (BIO.A.4.1.3 ) Transport Facilitation Student Name: Teacher Name: Jared

More information

Vorlesung Biophysik I - Molekulare Biophysik Kalbitzer/Kremer/Ziegler

Vorlesung Biophysik I - Molekulare Biophysik Kalbitzer/Kremer/Ziegler Vorlesung Biophysik I - Molekulare Biophysik Kalbitzer/Kremer/Ziegler 23.10. Zelle 30.10. Biologische Makromoleküle I 06.11. Biologische Makromoleküle II 13.11. Nukleinsäuren-Origami (DNA, RNA) 20.11.

More information

Lecture 6: Single nucleotide polymorphisms (SNPs) and Restriction Fragment Length Polymorphisms (RFLPs)

Lecture 6: Single nucleotide polymorphisms (SNPs) and Restriction Fragment Length Polymorphisms (RFLPs) Lecture 6: Single nucleotide polymorphisms (SNPs) and Restriction Fragment Length Polymorphisms (RFLPs) Single nucleotide polymorphisms or SNPs (pronounced "snips") are DNA sequence variations that occur

More information

MUTATION, DNA REPAIR AND CANCER

MUTATION, DNA REPAIR AND CANCER MUTATION, DNA REPAIR AND CANCER 1 Mutation A heritable change in the genetic material Essential to the continuity of life Source of variation for natural selection New mutations are more likely to be harmful

More information

Cis-and trans-isomer form of resvertrol

Cis-and trans-isomer form of resvertrol The Mechanism of Endothelium-Independent Relaxation Induced by the Wine Polyphenol Resveratrol in Human Internal Mammary Artery University of Belgrade Aleksandra Novakovic et.al University of Belgrade

More information

Antibody Function & Structure

Antibody Function & Structure Antibody Function & Structure Specifically bind to antigens in both the recognition phase (cellular receptors) and during the effector phase (synthesis and secretion) of humoral immunity Serology: the

More information

1 Mutation and Genetic Change

1 Mutation and Genetic Change CHAPTER 14 1 Mutation and Genetic Change SECTION Genes in Action KEY IDEAS As you read this section, keep these questions in mind: What is the origin of genetic differences among organisms? What kinds

More information

Knipholone anthrone from Kniphofia foliosa induces a rapid onset of necrotic cell death in cancer cells

Knipholone anthrone from Kniphofia foliosa induces a rapid onset of necrotic cell death in cancer cells Knipholone anthrone from Kniphofia foliosa induces a rapid onset of necrotic cell death in cancer cells Fitoterapia, 11/18/2010 Habtemariam S The present study examines the comparative cytotoxicity of

More information

Perioperative management of Dual Antiplatelet therapy post drug eluting stent-changing time

Perioperative management of Dual Antiplatelet therapy post drug eluting stent-changing time Perioperative management of Dual Antiplatelet therapy post drug eluting stent-changing time Robert Chilton DO, FACOI, FACC, FAHA Professor of Medicine University of Texas Health Science Center Director

More information

Integration and Coordination of the Human Body. Nervous System

Integration and Coordination of the Human Body. Nervous System I. General Info Integration and Coordination of the Human Body A. Both the and system are responsible for maintaining 1. Homeostasis is the process by which organisms keep internal conditions despite changes

More information

Graduate and Postdoctoral Affairs School of Biomedical Sciences College of Medicine. Graduate Certificate. Metabolic & Nutritional Medicine

Graduate and Postdoctoral Affairs School of Biomedical Sciences College of Medicine. Graduate Certificate. Metabolic & Nutritional Medicine Graduate and Postdoctoral Affairs School of Biomedical Sciences College of Medicine Graduate Certificate in Metabolic & Nutritional Medicine Graduate Certificate Metabolic & Nutritional Medicine Purpose

More information

Recombinant DNA and Biotechnology

Recombinant DNA and Biotechnology Recombinant DNA and Biotechnology Chapter 18 Lecture Objectives What Is Recombinant DNA? How Are New Genes Inserted into Cells? What Sources of DNA Are Used in Cloning? What Other Tools Are Used to Study

More information

Algorithms in Computational Biology (236522) spring 2007 Lecture #1

Algorithms in Computational Biology (236522) spring 2007 Lecture #1 Algorithms in Computational Biology (236522) spring 2007 Lecture #1 Lecturer: Shlomo Moran, Taub 639, tel 4363 Office hours: Tuesday 11:00-12:00/by appointment TA: Ilan Gronau, Taub 700, tel 4894 Office

More information

Neurotrophic factors and Their receptors

Neurotrophic factors and Their receptors Neurotrophic factors and Their receptors Huang Shu-Hong Institute of neurobiology 1 For decades, scientists believed that brain cells of the central nervous system could not regrow following damage due

More information

Copyright Mark Brandt, Ph.D. 46

Copyright Mark Brandt, Ph.D. 46 The Urea Cycle As has been mentioned, ammonium is toxic, and even small amounts will damage the nervous system. Genetic disorders in ammonium metabolism result in avoidance of high-protein foods and in

More information

Visualizing Cell Processes

Visualizing Cell Processes Visualizing Cell Processes A Series of Five Programs produced by BioMEDIA ASSOCIATES Content Guide for Program 3 Photosynthesis and Cellular Respiration Copyright 2001, BioMEDIA ASSOCIATES www.ebiomedia.com

More information

DNA Replication & Protein Synthesis. This isn t a baaaaaaaddd chapter!!!

DNA Replication & Protein Synthesis. This isn t a baaaaaaaddd chapter!!! DNA Replication & Protein Synthesis This isn t a baaaaaaaddd chapter!!! The Discovery of DNA s Structure Watson and Crick s discovery of DNA s structure was based on almost fifty years of research by other

More information

Introduction to Psychology, 7th Edition, Rod Plotnik Module 3: Brain s Building Blocks. Module 3. Brain s Building Blocks

Introduction to Psychology, 7th Edition, Rod Plotnik Module 3: Brain s Building Blocks. Module 3. Brain s Building Blocks Module 3 Brain s Building Blocks Structure of the Brain Genes chains of chemicals that are arranged like rungs on a twisting ladder there are about 100,000 genes that contain chemical instructions that

More information

Bottlenecks in Clinical Source Material Acquisition. Aby J. Mathew, PhD May 5, 2009 ISCT Annual Meeting San Diego, CA amathew@biolifesolutions.

Bottlenecks in Clinical Source Material Acquisition. Aby J. Mathew, PhD May 5, 2009 ISCT Annual Meeting San Diego, CA amathew@biolifesolutions. Bottlenecks in Clinical Source Material Acquisition Aby J. Mathew, PhD May 5, 2009 ISCT Annual Meeting San Diego, CA amathew@biolifesolutions.com Biopreservation What s the issue? Biopreservation considerations

More information

How many of you have checked out the web site on protein-dna interactions?

How many of you have checked out the web site on protein-dna interactions? How many of you have checked out the web site on protein-dna interactions? Example of an approximately 40,000 probe spotted oligo microarray with enlarged inset to show detail. Find and be ready to discuss

More information

3) There are different types of extracellular signaling molecules. 4) most signaling molecules are secreted by exocytosis

3) There are different types of extracellular signaling molecules. 4) most signaling molecules are secreted by exocytosis XIV) Signaling. A) The need for Signaling in multicellular organisms B) yeast need to signal to respond to various factors C) Extracellular signaling molecules bind to receptors 1) most bind to receptors

More information

BIOPHYSICS OF NERVE CELLS & NETWORKS

BIOPHYSICS OF NERVE CELLS & NETWORKS UNIVERSITY OF LONDON MSci EXAMINATION May 2007 for Internal Students of Imperial College of Science, Technology and Medicine This paper is also taken for the relevant Examination for the Associateship

More information

Introduction to Flow Cytometry

Introduction to Flow Cytometry Outline Introduction to Flow Cytometry Basic Concept of Flow Cytometry Introduction to Instrument Subsystems Daisy Kuo Assistant Product Manager E-mail: daisy_kuo@bd.com BDBiosciences Application Examples

More information

AP BIOLOGY CHAPTER 7 Cellular Respiration Outline

AP BIOLOGY CHAPTER 7 Cellular Respiration Outline AP BIOLOGY CHAPTER 7 Cellular Respiration Outline I. How cells get energy. A. Cellular Respiration 1. Cellular respiration includes the various metabolic pathways that break down carbohydrates and other

More information

GENE REGULATION. Teacher Packet

GENE REGULATION. Teacher Packet AP * BIOLOGY GENE REGULATION Teacher Packet AP* is a trademark of the College Entrance Examination Board. The College Entrance Examination Board was not involved in the production of this material. Pictures

More information