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Chapter 11 - Glycolysis 111 Glycolysis Is a Ubiquitous

Chapter 11 - Glycolysis 111 Glycolysis Is a Ubiquitous
Chapter 11 - Glycolysis 111 Glycolysis Is a Ubiquitous

Prentice Hall c2002Chapter 111Chapter 11 -Glycolysis

?For centuries, bakeries and breweries have exploited the

conversion of glucose to ethanol and CO 2by glycolysis in yeast

Prentice Hall c2002Chapter 112

11.1Glycolysis Is a Ubiquitous Pathway

Converts: 1 glucose

2 pyruvate

?Pyruvate can be further metabolized to:

(1) Lactate or ethanol (anaerobic) (2) Acetyl CoA (aerobic) ?Acetyl CoA is further oxidized to CO 2and H 2O via the citric acid cycle

?Much more ATP is generated from the citric acid cycle than from glycolysis

Prentice Hall c2002

Chapter 11

3

Fig 11.1

?Catabolism of glucose via glycolysis and the citric acid cycle

(continued next slide)

Prentice Hall c2002

Chapter 11

4

Fig 11.1 (continued)

Prentice Hall c2002Chapter 115Net reaction of glycolysis ?Two molecules of ATP are produced

?Two molecules of NAD +are reduced to NADH

Glucose + 2 ADP + 2 NAD + + 2 P i

2 Pyruvate + 2 ATP + 2 NADH + 2 H ++ 2 H 2O

Prentice Hall c2002Chapter 116

Table 11.1

Prentice Hall c2002Chapter 117Glycolysis (10 reactions) can be

divided into two stages

?Hexose stage: 2 ATP are consumed per glucose ?Triose stage: 4 ATP are produced per glucose

Net: 2 ATP produced per glucose

Prentice Hall c2002Chapter 118

Fig 11.2

?Glycolysis (next 4 slides)

Prentice Hall c2002Chapter 119Phosphofructokinase-1

Prentice Hall c2002Chapter 1110

Prentice Hall c2002

Chapter 11

11

Prentice Hall c2002

Chapter 11

12

11.2Glycolysis Has 10 Enzyme-Catalyzed Steps

?Each chemical reaction prepares a substrate for the next step in the process ?A hexose is cleaved to two trioses

?Interconversion of the trioses allows both to be further metabolized via glycolytic enzymes ?ATP is both consumed and produced in glycolysis

Prentice Hall c2002

Chapter 11

13

1.Hexokinase

?Transfers the γ-phosphoryl of ATP to glucose C-6 oxygen to generate glucose 6-phosphate (G6P ) ?Mechanism: attack of C-6 hydroxyl oxygen of glucose on the γ-phosphorous of MgATP 2-displacing MgADP -?Four kinases in glycolysis: steps 1,3,7, and 10?All four kinases require Mg 2+and have a similar mechanism

Prentice Hall c2002Chapter 1114

Fig 11.3Hexokinase reaction

Prentice Hall c2002Chapter 1115

Properties of hexokinases ?Broad substrate specificity -hexokinases can phosphorylate glucose, mannose and fructose ?Yeast hexokinase undergoes an induced-fit conformational change when glucose binds ?Conformational change helps prevent hydrolysis of ATP to ADP and P i (Fig 6.13)

Prentice Hall c2002

Chapter 11

16

Isozymes of hexokinase

?Isozymes -multiple forms of hexokinase occur in mammalian tissues and yeast

?Hexokinases I, II, III are active at normal glucose concentrations (K m values ~10-6to 10-4M)?Hexokinase IV (Glucokinase , K m ~10-2M) is

active at higher glucose levels, allows the liver to respond to large increases in blood glucose

Prentice Hall c2002

Chapter 11

17

2. Glucose 6-Phosphate Isomerase

?Converts glucose 6-phosphate (G6P ) (an

aldose) to fructose 6-phosphate (F6P ) (a ketose)?Enzyme preferentially binds the α-anomer of G6P (converts to open chain form in the active site) ?Enzyme is highly stereospecific for G6P and F6P ?Isomerase reaction is near-equilibrium in cells

Prentice Hall c2002Chapter 1118

Fig 11.4 Conversion of G6P to F6P

Prentice Hall c2002

Chapter 11

19

3.Phosphofructokinase-1 (PFK-1)

?Catalyzes transfer of a phosphoryl group from ATP to the C-1 hydroxyl group of F6P to form fructose 1,6-bis phosphate (F1,6BP )

?PFK-1 is metabolically irreversible and a critical regulatory point for glycolysis in most cells (PFK-1 is the first committed step of glycolysis)?A second phosphofructokinase (PFK-2)

synthesizes fructose 2,6-bis phosphate (F2,6BP )

Prentice Hall c2002

Chapter 1120

PFK-1 Reaction

Prentice Hall c2002

Chapter 11

21

4.Aldolase

?Aldolase cleaves the hexose F1,6BP into two triose phosphates:glyceraldehyde 3-phosphate (GAP ) and dihydroxyacetone phosphate (DHAP )

?Reaction is near-equilibrium, not a control point ?Mechanism is common for cleaving C-C bonds in biological systems (and C-C bond formation in the reverse direction)

Prentice Hall c2002

Chapter 1122

Aldolase Reaction

Prentice Hall c2002Chapter 1123Fig 11.5 Mechanism of aldolases

(continued next slide)

Prentice Hall c2002Chapter 1124

Fig 11.5 (continued)

Prentice Hall c2002

Chapter 11

25

5.Triose Phosphate Isomerase (TPI)?Conversion of DHAP into glyceraldehyde 3-phosphate (GAP)

?Reaction is very fast (diffusion controlled), and only the D-isomer of GAP is formed

?Radioisotopic tracer studies show:

One GAP molecule: C1,2,3 from Glucose C4,5,6

Second GAP: C1,2,3 from Glucose C3,2,1

Prentice Hall c2002

Chapter 11

26

Reaction of Triose

phosphate isomerase

Prentice Hall c2002Chapter 1127Fig 11.6 Fate of carbon atoms from

hexose stage to triose stage

Prentice Hall c2002Chapter 1128

6.Glyceraldehyde 3-Phosphate

Dehydrogenase (GAPDH)

?Conversion of GAP to

1,3-bis phosphoglycerate (1,3BPG )?Molecule of NAD +is reduced to NADH

?Oxidation of the aldehyde group of GAP

proceeds with large negative free-energy change

Prentice Hall c2002Chapter 1129Conservation of oxidative energy

?Energy from oxidation of GAP aldehyde is conserved in acid-anhydride linkage of 1,3BPG ?Next step of glycolysis uses the high-energy phosphate of 1,3BPG to form ATP from ADP ?Mechanism of GAPDH shows how an energy-rich compound forms in an oxidation reaction

Prentice Hall c2002Chapter 1130

Reaction of GAPDH: GAP converted to 1,3BPG

Prentice Hall c2002Chapter 1131Fig 11.7?Mechanism of GAPDH (3 slides)

Prentice Hall c2002Chapter 1132

Fig 11.7 (continued)

(2)

(4)

Prentice Hall c2002Chapter 1133Fig 11.7 (continued)

Prentice Hall c2002Chapter 1134

Box 11.2 Arsenate (AsO 43-) poisoning

?Arsenate can replace P i as a substrate for G3PDH ?Arseno analog which forms is unstable

Prentice Hall c2002

Chapter 11

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7.Phosphoglycerate Kinase (PGK)

?Transfer of phosphoryl group from the energy-rich mixed anhydride 1,3BPG to ADP yields ATP and 3-phosphoglycerate (3PG )?Substrate-level phosphorylation -Steps 6 and 7 couple oxidation of an aldehyde to a carboxylic acid with the phosphorylation of ADP to ATP

Prentice Hall c2002

Chapter 11

36

Phosphoglycerate kinase reaction

Prentice Hall c2002

Chapter 11

37

8.Phosphoglycerate Mutase

?Catalyzes transfer of a phosphoryl group from one part of a substrate molecule to another ?Reaction occurs without input of ATP energy ?Mechanism requires 2 phosphoryl-group transfer steps

?Enzymes from animal muscle and yeast have a different mechanism than does plant enzyme

Prentice Hall c2002

Chapter 11

38

Phosphoglycerate mutase reaction

Prentice Hall c2002Chapter 1139Fig 11.8Phosphoglycerate mutase

mechanism: animals and yeast

(continued)

Prentice Hall c2002Chapter 1140

Fig 11.8 (continued)

(1)

Prentice Hall c2002Chapter 1141Fig 11.8 (continued)

Prentice Hall c2002Chapter 1142

9.Enolase: 2PG to PEP

?3-Phosphoglycerate (3PG ) is dehydrated to phosphoenolpyruvate (PEP )

?Elimination of water from C-2 and C-3 yields the enol-phosphate PEP

?PEP has a very high phosphoryl group transfer potential because it exists in its unstable enol form

Prentice Hall c2002

Chapter 11

43

Enolase reaction

Prentice Hall c2002

Chapter 11

44

10.Pyruvate Kinase (PK)

?Catalyzes a substrate-level phosphorylation ?Metabolically irreversible reaction

?Regulation both by allosteric modulators and by covalent modification

?Pyruvate kinase gene can be regulated by various hormones and nutrients

PEP + ADP

Pyruvate + ATP

Prentice Hall c2002Chapter 1145Fig 11.9

?Pyruvate kinase reaction

Prentice Hall c2002Chapter 1146

11.3The Fate of Pyruvate

1. Aerobic conditions: oxidized to acetyl CoA which enters the citric acid cycle for further oxidation

2. Anaerobic conditions (microorganisms): conversion to ethanol

3. Anaerobic conditions (muscles, red blood cells): conversion to lactate

Prentice Hall c2002Chapter 1147Fig 11.10

?Three major fates of pyruvate

Prentice Hall c2002Chapter 1148

A. Metabolism of Pyruvate to Ethanol

(yeast -anaerobic)

?Two reactions required: (1) Pyruvate carboxylase (2) Alcohol dehydrogenase Pyruvate

Acetaldehyde Ethanol

(1)

(2)

Prentice Hall c2002Chapter 1149Fig 11.11

?Anaerobic conversion of

pyruvate to ethanol (yeast)

Prentice Hall c2002Chapter 1150

Fig 11.11 (cont)

Prentice Hall c2002Chapter 1151

B. Reduction of Pyruvate to Lactate

(muscles -anaerobic)

?Muscles lack pyruvate dehydrogenase and cannot produce ethanol from pyruvate ?Muscle lactate dehydrogenase converts pyruvate to lactate

?This reaction regenerates NAD +for use by GAPDH in glycolysis

Prentice Hall c2002

Chapter 11

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Recycling of lactate

?Lactate formed in skeletal muscles during exercise is transported to the liver ?Liver lactate dehydrogenase can reconvert lactate to pyruvate

?Lactic acidosis can result from insufficient

oxygen (an increase in lactic acid and decrease in blood pH)

Prentice Hall c2002Chapter 1153Reduction of pyruvate to lactate

Prentice Hall c2002Chapter 1154

Overall reactions for glucose

degradation to lactate

?Two ATP per molecule glucose consumed ?No oxygen is required Glucose + 2 P i 2-+ 2 ADP 3-2 Lactate -+ 2 ATP 4-+ 2 H 2O

Prentice Hall c2002

Chapter 11

55

11.4Free-Energy Changes in Glycolysis

?Actual free-energy changes (?G) large only for:

#1 (hexokinase)

#3 (phosphofructokinase)#10 (pyruvate kinase)?These steps are metabolically irreversible, and these enzymes are regulated

??G for all other steps are close to zero (they are near-equilibrium in cells)

Prentice Hall c2002Chapter 1156

Fig 11.12 Cumulative standard and actual free energy changes for the reactions of glycolysis

Prentice Hall c2002

Chapter 11

57

11.5Regulation of Glycolysis

1. When ATP is needed, glycolysis is activated ?AMP and fructose 2,6-bis phosphate (F2,6BP) relieve the inhibition of PFK-1 by ATP

2. When ATP levels are sufficient,glycolysis activity decreases ?PFK-1 is inhibited by ATP and citrate

?Hexokinase inhibited by excess glucose 6-phosphate

Prentice Hall c2002

Chapter 11

58

Fig 11.13 Metabolic regulation of glycolysis

(continued)

Prentice Hall c2002Chapter 1159Fig 11.13 (continued)

Prentice Hall c2002Chapter 1160

A. Regulation of Hexose Transporters

?Glucose enters mammalian cells by passive

transport down a concentration gradient from blood to cells

?GLUT is a family of six passive hexose transporters ?Glucose uptake into skeletal and heart muscle and adipocytes by GLUT 4 is stimulated by insulin ?Other GLUT transporters mediate glucose transport in and out of cells in the absence of insulin

Prentice Hall c2002Chapter 1161Fig 11.14 Regulation of glucose

transport by insulin

Prentice Hall c2002Chapter 1162

B. Regulation of Hexokinase

?Hexokinase reaction is metabolically irreversible ?G6P (product) levels increase when glycolysis is inhibited at sites further along in the pathway ?G6P inhibits hexokinase isozymes I, II and III ?Glucokinase forms G6P in the liver (for glycogen synthesis) when glucose is abundant (activity is modulated by fructose phosphates and a regulatory protein)

Prentice Hall c2002Chapter 1163Fig 11.15 Effects of a regulatory protein

on glucokinase kinetics

?Addition of a

regulatory protein raises apparent K m for glucose (inhibits glucokinase)

Prentice Hall c2002Chapter 1164

Box 11.3 Glucose 6-Phosphate Has a

Pivotal Metabolic Role in Liver

Prentice Hall c2002

Chapter 11

65

C. Regulation of Phosphofructokinase-1

?ATP is a substrate and an allosteric inhibitor of PFK-1?AMP allosterically activates PFK-1 by relieving the ATP inhibition (ADP is also an activator in mammalian systems)

?Changes in AMP and ADP concentrations can control the flux through PFK-1

?Elevated levels of citrate (indicate ample substrates for citric acid cycle) also inhibit PFK-1

Prentice Hall c2002

Chapter 11

66

Fig 11.16 Regulation of PFK-1 by ATP and AMP

?AMP relieves ATP inhibition of PFK-1

Prentice Hall c2002Chapter 1167Regulation of PFK-1 by Fructose 2,6-bis phosphate (F2,6BP)

?F2,6BP is formed from F6P by the enzyme phosphofructokinase-2 (PFK-2)

?Fig 11.17β-D-Fructose 2,6-bis phosphate

Prentice Hall c2002Chapter 1168

Formation and hydrolysis of F2,6BP

Prentice Hall c2002Chapter 1169Fig. 11.18

?Effect of glucagon on liver glycolysis

Prentice Hall c2002Chapter 1170

D. Regulation of Pyruvate Kinase (PK)

?Four PK isozymes exist in mammalian tissues ?PK is allosterically activated by F1,6BP,inhibited by ATP

?Glucagon stimulates protein kinase A which phosphorylates PK converting it to a less active form (liver and intestinal cells)

Prentice Hall c2002Chapter 1171Fig 11.19 Initial velocity curves of

pyruvate kinase

Prentice Hall c2002Chapter 1172

Fig 11.20Pyruvate kinase with F1,6BP

?Activator F1,6BP (red)?Active site is in large central domain

Prentice Hall c2002

Chapter 11

73

E. The Pasteur Effect

?Under anaerobic conditions the conversion of glucose to pyruvate is much higher than under aerobic conditions (yeast cells produce more ethanol and muscle cells accumulate lactate)?The Pasteur Effect is the slowing of glycolysis in the presence of oxygen

?More ATP is produced under aerobic conditions than under anaerobic conditions, therefore less glucose is consumed aerobically

Prentice Hall c2002

Chapter 11

74

11.6Other Sugars Can Enter Glycolysis

?Glucose is the main metabolic fuel in most organisms ?Other sugars convert to glycolytic intermediates ?Fructose and sucrose (contains fructose) are major sweeteners in many foods and beverages ?Galactose from milk lactose (a disaccharide) ?Mannose from dietary polysaccharides,glycoproteins

Prentice Hall c2002Chapter 1175A. Fructose Is Converted to Glyceraldehyde 3-Phosphate

Fig 11.21

(continued)

Prentice Hall c2002Chapter 1176

Fig 11.21 (continued)

Prentice Hall c2002Chapter 1177B.Galactose is Converted to

Glucose 1-Phosphate

Fig 11.22 (continued next slide)

Prentice Hall c2002Chapter 1178

Fig 11.22 (continued)

Prentice Hall c2002Chapter 1179C. Mannose is Converted to Fructose 6-Phosphate

Prentice Hall c2002Chapter 1180

11.7Formation of 2,3-Bis phosphoglycerate

in Red Blood Cells

?2,3-Bis phosphoglycerate (2,3BPG)allosterically regulates hemoglobin oxygenation (red blood cells)?Erythrocytes contain bis phosphoglycerate mutase which forms 2,3BPG from 1,3BPG

?In red blood cells about 20% of the glycolytic flux is diverted for the production of the important oxygen regulator 2,3BPG

汇付和托收有答案

汇付和托收 一、翻译并解释下列名词 1.Remittance 2.T/T 3.M/T 4.Collection 5.Collection 6.D/P 7.D/A 8.D/P.T/R 二、问答题 1.简述汇付结算方式的特性。 2.简述托收业务的种类。 3.比较即期付款交单、远期付款交单、承兑交单的异同。 4.为什么在出口业务中不宜轻易按远期D/P方式成交 5.D/P at30 days after sight和D/A at 30days after sight有何区别各有何风险} 6.简述托收方式的特点。 三、单项选择题 1.托收方式收取货款采用的是( )。 A.顺汇方法B.逆汇方法C.反推方法D.信汇方法 2.在托收方式下,卖方委托银行收取货款,使用的汇票是( )。 A.商业汇票,属于商业信用 B.银行汇票,属于银行信用

C.商业汇票,属于银行信用 D.银行汇票,属于商业信用 】 3.( )方式只适用于远期汇票的托收。 A.付款交单B.承兑交单C.信托交单D.顺汇方法 4.在托收方式下,对出口商来说,下面哪一种选择风险最小( )。 A.D/P 90天B.D/A 60天C.D/P 30天D.D/A 30天 5.我们所说的托收业务属于商业信用是因为( )。 A.没有银行参与B.出票人开立的汇票是银行汇票C.银行不承担保证付款的义务D.以上都对 6.使用托收方式时,托收行和代收行在收回货款方面( )。 A.没有责任B.承担部分责任C.有责任D.视合约而定 7.D/P和D/A支付方式下,就卖方风险而言,哪个正确( )。 - A.D/A>D/P B.D/P>D/A C.D/A=D/P D.不确定 8.合同中规定采用D/P 30天,如果托收日为8月l it,寄单邮程为7天,则此业务的提示日、承兑日、付款日分别为( )。 A.8月8 13/8月8日/9月7 日B.8月1 日/9月6 日/8月1 日C.9月6 13/8月1日/8月1 日D.8月1 日/9月6 日/9月6 日9.承兑交单的起算日应为( )。

郭著章《英汉互译实用教程》(第4版)教材配套题库-第5章 翻译常用的八种技巧【圣才出品】

第5章翻译常用的八种技巧 5.1 重译法 一、英译汉 1. It may seem strange to put into the same packet an industrial revolution and two political revolutions. But the fact is that they were all social revolutions. 【译文】把一场工业革命同两次政治革命归作一类似乎有点奇怪,但事实上这三次革命都是社会革命。 2. Two things are outstanding in the creation of the English system of canals, and they characterise all the Industrial Revolution. 【译文】在修建英国的运河网的过程中,有两点是非常突出的,而这两点也正是整个工业革命的特点。 3. The canals were arteries of communication: they were not made to carry pleasure boats, but barges. 【译文】运河是交通的动脉,开运河不是为了走游艇,而是为了通行驳船。 4. James Brindley was a pioneer in the art of building canals or, as it was then called, ‘navigation’. 【译文】布林德雷是开凿运河的先驱者,当时人们把开凿运河叫做navigation。

5. Several factors accounted for this extraordinary achievement. One was the expansion into the west. Another was the application of machinery to farming. 【译文】取得这一特殊成就有几方面的原因:第一个原因是向西部的扩展,第二个原因是机器在农业上的应用。 6. …for the establishment of agricultural and industrial colleges. These were to serve both as educational institutions and as centers for research in scientific farming. 【译文】……以便建立农业和工业学院。这些学校一方面是教育机关,另一方面,也是农业科学的研究中心。 7. …This is nothing li ke back home in Colorado. We have rains there, too. Thunderstorms in spring and summer... 【译文】……这跟家乡科罗拉多的情况迥然不同。科罗拉多也下雨。春夏两季雷雨交加……【解析】原文第二句用there代替in Colorado,而译文重复“科罗拉多”。 8. The world watches. The world listens. The world waits to see what we will do. 【译文】全世界在注视着。全世界在倾听着。全世界在等待着看我们将做些什么。 9. As you said in your toast, the Chinese people are a great people, the American people are a great people.

国际贸易实务课后答案详解-第十一章--汇付和托收

第十一章汇付和托收 一、思考题 1.试写出T/T、M/T、D/D、D/P、D/A、T/R、O/A的英文全文、中文译名,并分别简述其基本含义。 答:(1)电汇(telegraphic transfer,T/T)是由汇款人委托汇出行用电报、电传、环球银行间金融电讯网络(SWIFT)等电讯手段发出付款委托通知书给收款人所在地的汇入行,委托它将款项解付给指定的收款人的一种汇付方式。 (2)信汇(mail transfer,M/T)是指汇出行应汇款人的申请,以信汇委托书或支付通知书作为结算工具,通过邮政航空信件方式寄发给汇入行的一种汇付方式。 (3)票汇(remittance by banker’s demand draft,D/D)是指汇出行应汇款人的申请,以票据作为结算工具,开立以其代理行或其他往来银行为付款人的银行即期汇票,列明收款人名称、金额等,交由汇款人自行寄交给收款人,凭票向付款行取款的一种汇付方式。 (4)付款交单(documents against payment,D/P)是指出口人的交单须以进口人的付款为条件,即出口人将汇票连同货运单据交给银行托收时,指示银行只有在进口人付清货款时才能交出货运单据。 (5)承兑交单(documents against acceptance,简称D/A)是指出口人的交单以进口人的承兑为条件,进口人承兑汇票后,即可向银行取得货运单据,待汇票到期日才付款。 (6)信托收据(trust receipt,T/R)是进口人借单时提供的一种书面信用担保文件,用以表示出据人愿意以代收银行的受托人身份代为提货、报美、存仓、保险、出售,同时承认货物的所有权仍属银行。 (7)货到付款也称“赊账交易”(Open Account transaction,简称O/A),是指在签订合同后,出口人先将货物发出,进口人收到货物后立即或在约定的一段时间后通过银行付款。 2.“在国际货物买卖中,对买卖双方来说,汇付是一种最安全的支付方式”,这种认识是否正确?为什么? 答:这种认识不正确。理由如下: 汇付方式属于商业信用性质。银行仅凭汇款人的指示转移相关款项,并不负责单据的传递,更不承担任何付款或担保责任。使用汇付方式完全取决于买卖双方中的一方对另一方的信任,并在此基础上向对方提供信用和进行资金融通。 3.为什么说汇付和托收分属顺汇和逆汇,但又都属商业信用性质? 答:(1)汇付和托收分属顺汇和逆汇的原因 汇付又称汇款,是指订立商务合同后,进口人(汇款人)通过银行向出口人(收款人)汇寄款项的做法。由于汇款业务中使用的结算工具(委托通知、票据)的传递方向与资金的流向相同,所以汇付方式属于顺汇。 托收是指由接到委托指示的银行处理金融单据和/或商业单据以便取得承兑或付款,或凭承兑或付款交出商业单据,或凭其他条件交出单据。由于托收业务中使用的结算工具(托收指示书和汇票)的传送方向与资金的流动方向相反,所以托收方式属于逆汇。 (2)汇付和托收都属于商业信用的原因 因为汇付结算货款的过程中和托收业务中,银行都只是提供服务而不提供信用,所以汇付和托收都属于商业信用性质。 4.试比较凭单付汇与一般汇付和跟单托收的异同。

实用英汉翻译教程答案

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