Liquid Chromatography Mass Spectrometry 3rd Edition

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Dedication
I would like to ded icate thi s 1"ll.1I1k to my mother. Lott ie Dud ek . who was bo rn on Novembe r 11, 1918.
Through the years, my mo th er raised h er child ren, maintained a In,vi n~ marr iage. cared fl'f h er a,ging
paren ts, and worked 40 hou rs per wee k . In 1990s parla nce. soc iety would desc ribe such ,I person as a
"liberated wom an" o r "supermom." ] would like to acknowledge rbar mv mother was 11 "supenruuu"
20 years before th e word was fashio nable. A son cannot repay a mot her. My hope is that "I lov c' yllll
and th ank you" will suffice.
Contents

Preface •••• •• •••• • •• • • • • • • • . • • • • • • • • • ••• •••• • •••••• • • • • . • xiii

Cell Biology ••• • •. . . .... . . . . 1
I. Nuc lear struc tures I
II. Cytoplasm 2
III. Cy topb smic struct ures 2
IV. Cytoskeleton 6
v. c-umembrane 7
VI. Cell cycle 10
VII. Apo prosis 11
VIII. Ce ll incl us io ns 1)
IX. Selected phorormcrog raphs 13

2 Epitheliu m •••• • • • • • • • • •. . . . 19
I.Introduction 19
II.Classificat ion 19
Ill.Polarity 19
IV. Cl in ical cons iderations 22
V. Sell'Cted phot .>lnicrognl.plu 22

3 C onnective Tissue ...................... ..... ...... ..... 25

I. Inrnxl uction 25
II. Ground substance 25
III. Fi~ 15
IV. Ce lls 26
V. C linical cons iderations 28
VI. Selected phoromicrograpN 30

4 Car tilage •••••• •• • • . . . • • • • • • • . . . . .•••• • •• •• • • • • • ••••••• 31

I. Int roduction J I
II. Gro und substance 31
III. Fih: rs 31
IV. Cd ls 31
V. Blond vessels and nerves 31
VI. Chondrogenesis 31
VII. Hormonal influence 32
VIII. Repair 32

5 Bone. . . . . . . . . .... . . ... . . . ... ..... . . .. . . . . . . . . . ... . . . . 33

I. Introduction 33
II. G round substance 33
III. Fibers 33
IV. Ce lls 33
V. Blood vessels and nerves H

ril
vllt Contents

VI. Osteogenesis H
VII. flo"" r"p"ir 35
VIII. HOrllw l\al influence 35
IX. C lin ica l conside rat ion s 36
X. C 'lrtilaJ.:e and ho ne cnm pnnson 37
XI. Selected photomicrographs 38

6 Muscle .•••.•• .. ....... .. ...... ... ......... ... ..... 41

I. Skd"tal muscle 41
II. C ardiac muscle 46
III. Smooth muscle 46
IV. Co mparisons and con trasts 47
V. Selected photomicrographs 48

7 Nervous TIssue . . . . . . . 51
I. Th", IWUro" 51
II. Neurogl ial cells '52
III. Th e blood-brain barrier 55
IV. TIle blood.. . C SF harrier 51
V. Ner ve degene rat ion and regenerati on 55
VI. Clinical cons iderations 56
VII. Selec ted photomicrographs 57

8 He art and Blood Vessels •..

I. Heart lnvcrs 63
......... .... ....... .. .... .. 63
II. Cont ract ion of cardiac mvocv rcs 6'5
III. C onduct ion system 65
IV. Neural r",gu!ali",' "fhe"rt rare 6 7
V. Enzyme levels in myoc ard ial infarction 68
VI. Blood vessels 68
VII. Func t io ns of endothel ium 70
VIII. Blood flow 70
IX. Types of circulation 71
X. Select ed photomicrographs 71

9 Blood . 78
I. rl"~llIa 78
II . R"J bluuJ cells 78
III. Hemoglohin 79
IV. Blood gas exchan ge 79
V. Whi te blood cel l> 81
VI. H ~'ps.·rSt"mit ivity r",,,ct iollS 83
VII. Platele ts 84
VIII. Hemo stasis 84
IX. Selec ted phot omicrographs 86

10 Thymus . 95
I. Thy m ic co rte x 95
II. Thymic medulla 9'5
III. Maturation of T ce lls 95
IV. Rlood -·thymus harrier 95
V. Involution of th e thy mus 97
VI. Thyme ctomy 9 7
 COntents Ix

1.1 Lymph Nod e . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 98

I. o..le r co rtex 98
II. Inne r co rte x 98
III. Medulla 9M
IV. Flow oflYlllph 98
V. Flow of blood 98
VI. Cluneal cons ide ration 99
VII. Selected photomicrographs 100
12 Spleen .... .. . .. . .. . . . . . . ••. . . . . . . . ... . . . . . . . . . . . . . . . . 10 1
I. Whit t' pulp 101
II. M;u-ginal woe 101
III. ReJ pulp 101
IV. Blood tlow 10I
V. C lin ical co nsideratums !OI

13 St omach .. . • • .... . ... . . . • • . ..• • •• • • • •. . • • • • • • • • • . • . . • . 103

I. Mucosa 103
II. G amic glanJs 103
III. G astr ic emply inK 105
IV. Ro:pair (rege ne rat ion) 105
V. Cl in ical consiJrrat itlll.\. 105
14 Small I ntest ine • • • •• •• •• • •..• . . .• • • • • • • • • • • • •• • • • • • . •. .. 106
I.Mucosa 106
II.[n test ma] xlan,h (c rypts of Lleberku hn) 107
III.Gut-associated Iympha lic tis.suc 107
IV. Repair (rcgcncrunon) 10 7
V. C lin ical cn nsidl'l1lfions 107
15 Large Intestine (Colon) •••• • •• ••• .• . . . . .. . . . . . . . .. • • • •.... III
I. Anal canal I II
II. Mucosa III
III. Intestin al glanJ.~ III
IV. Gue.assooared lymphat ic tissue III
V. Repa ir Iregcne ranon) III
VI. C linica l co nsidera uore III

16 L iver •• ••• ••• • • • . . . . .. . . . . . . . . ...... • . . . . . . . .. ... . . .. 116

I. Hcpa tocvrcs 116
II. Kupffer cells 117
III. Lipocvtes (fat -stor ing cells; Ito cells) 117
IV. C lassic liver lobu le 117
V. U vcr acinu s 118
VI. l ymph 118
VII. Repair (r~nt'r.lf ion ) 119
VIII. C hn lcal co nstderanons 119
IX. Se lected pho tomic rographs 110

17 Exocrin e Pa ncreas and Is lets of Langerhans . . . . . . . . . . . . . . . . . . . . 123

I. Exoc rine pancreas 113
II. lslers of Langcrhans 123
Ill. Insulin recep tor and signal transduct ion I Z3
IV. C lin ica l co nside rations 126
V. Se lected phohlm icrographs 128
It Contents

1 8 Respirat ory System .. . .. • •. •.. . . . • •• • • •• . . . . • • •••• • •. • • • • 130

I. Cordocnon port ion 130
II. R~piratory po rtion I}I
III. Surfacla nt 13I
IV. ComponenlS ol rhr bl<lu J-alr barrier 131
V. Repair (regeneration) 131
VI. Chrucal consideranoes I} I
VII. Selected pho lom icnwarhs I}l

19 U rina ry System • •• • • • . • . ..••• • • •. . . .. •. •. . . . .. . . ...... . 136

I. Renal [cnnifcrccs] t uhules 136
II. Nephrons 136
III. The collec tm g duet 137
IV. Filtra tion barncr 137
V. Th e juxtaglomerular complex 138
VI. H isw physiology 138
VII. Th.. co unt ercurrenr multiplier system 138
VIII. The cou ntercurrent exc han}:a syste m 139
IX. Glomerular filtrat ion rate 139
x. C learance 140
XI. C lin ical coruderenons 141
XII. Selected phm omicnwarhs 142

20 Hypophysis • • . . . . • . • • •• • • •• • • • . ••• • ••.•.• • . • . . . . . . ..... 147

I. Th e aden ohypophy sis 147
II. Hoenoeal secrenon 147
III. The neulUhYPI.Jrhysis 148

21 T hy roid . . . . .. . . . . . . . . .. . . . .• . . . . .. . . . . . . . . . . . . . . . . . . • 150
I. Thyroid follicles I SO
II. Follic ular cel ls I SO
III. Fu nct ions ofTJ and T~ 150
IV. Pamfollicular ce lls 152
V. O in ical cons iderations 152
VI. Se lected phOlOmicl"Ofo(r.tl'hs I S4

22 Parathyroid . • • . . . . . . . • • • • • • • • . . • • • • • • • • . . • • • • • . . . . . . . .. 156
I. Chiefcells ! 56
II. O xvphil ce lls 156
III. Calcium h()me".,t,, ~i s I S6
IV. C limcal consideratio ns 156

23 Adrenal • • . . . •• • • • • •••• . . . .•• •• •• . . . . . . . .. . . . .••• ••• •• 158

I. Co rtex 158
II. Th e medulla 161
III. Selected phuto micnlb'rdrhs 165

24 Female Reproductive System 167

I. Adult ovary {cortex} 167
II. Corpus luteum 167
III. Uterine tubes 167
IV. Uterus 168
V. Ce rvix 171
 COntents III

VI. Va~ina 171

VII. Mammarv gland In
VIII. Selected rhOlomic~rh.s 173

25 Male Reproductive System .... ... . .. •. . . . .• ....... . . . . . . . . 178

I. ~m in ifcrous tubules J78
II. Srcnn iogenesis 178
Ill. Sperm morpholos:y 179
IV. Leydig ( interstitial) cells 179
V. Prostate gland 182
VI. C linical ccnstdera nons 182
VII. Se lected photomicrographs 183

26 Skin . . . . . . . . . . . . . . . . . . • • • • • • • • •• 190
I. Epidermis 190
II . Derm is 190
III. G lands 190
IV. Nerves 191
V. C lin ical co nsi de ra tions 192

27 Ear . .. . . .. . .. . • • • •..... ••••• . . . . . ... ..... . . . . ... ..... 19 3

I. Tympanic membrane (eard rum) 193
II. Membranous labyrinth 193
III . C linical consideration 195

28 Eye • . . . .• . . • . ... ••• • ••• . ..• . . . . . . . . . . . . . . . . . . . . . . . . . . 196

l. Cornea 196
II. limbus 196
III. Ret ina 196
IV. Visual rransducnon 196
V. C lin ica l consiJt. mrim15 199
VI. Se lected phowm icto,.:raphs 200

29 Cerebral Cortex • •• •• . . . . •• • . . . . . . . . . • . ..• . . . . . . .. . . .... 202

I. C yroa rch trec mre 202
II. Orgame at ton 202
III. Selected phoronucrographs 203

30 Cerebellar Correx ... • • • . ... . . . • • • • . . . . . . . . . . . . . . . . . . . . .. 205

I. Cytoarchi tect ure 20 5
II. Ce rebellar !<lomcrullis 208
III. C lin ical cousid eranous 208

Index . . . . • . • . • • • • . • . . . . . • • • • . • .. 209
Preface
The seco nd ed it ion of H igh~Yidd Hi5wlu.!:J h as bee n a pleasure to wri te. Fo llowing: rhe publicanon of
the first edition. I received man y co mme n ts from readers and revte w crs co ncern ing how we ll th e hook
me t th e stared goal of the High.YItId series, specifica lly, [ 0 pro vide uncom plicated , co nc ise coverage
of only those WpiC5 mat are extremel y relevant when one is preparing (or th e Uni ted Smre, Med ical
licensing Exam ination (USM LE). In preparing th e second edition, I have responded Itl rhc~ com -
ments.
This exercise h as led to th e developm en t of an un usua l type of h istology book . As you ma y kn ow, th e
questions on th e USMLE cross tradit ional co urse boundaries, makin g it d iffi cult to idcnt ify a ques-
tion th at is "stric tly h istology." Man y US MLE q uest ions fall inr o ca tegories such as hist opathologv,
h istop h ysio logy, and htstobiochem istry, To th is end, the second ed it ion of Hi~h-Yield Hi~ lUlu!D' rev iews
basic h istologic co ncepts and the n exte nd s th e d iscussion ro relevant areas in pathologv, physio k1{Y.
and biochemistry. This makes High~Y"aeld Hi5fOlogy a truly In tegrated review book that reflects th e way
histolem' is tested on the US MlE Step 1.
In eddmon, many students have commented m a t cell biolOJ.'Y and bhxl top ics have been well rep -
resented on the USMlE Step I . To th is end , I have expanded Chapter I. "C ell Biolow,'," and added
a chapte r (Chapter 9 , "Blood").
High-Yield HiswloK), 2nd edition; High-Yield EmbryololO . H igh~Yield Gross Anatomy. and Hil{h-Yield
Mulecular Biulogy comprise my co n tribution to th e High-Yield series. I would apprecia te any comments
or suggest ions co ncern ing any of these revie w honks, especially afte r you have taken the USMlE S rcp
l. You may con tact me at d [email protected] u. edu.

llil
 1.
Cell Biology

I. NUCLEAR STRUCTURES
A. N uclear envelope
1. The inn er membrane is associ ated with n net work of intermediate filament s
{lam in a A, B. C) ca lled the nuclear lamin a, wh ich plays a role in the re-assembly
of the n uclear env elope during teloph ase of m itosis. T h e oute r membrane is stud-
ded with ribosomes and is cont inuous with the rough endoplasmic ret icul um
(,ER).
2 . The inn er and outer membranes are separated by a perinuclea r cisterna .
3 . The nu clear pore complex consists of many different proteins arranged in cctag -
a na l sym metry with a centra l ch anne l. The nuclear pore co mplex allows pessage
of molecules betwee n the nucleus and cytoplasm (Table 1. 1).
B. Chromatin is double-helical DN A associated with histones and nonhistone proteins.
1. H eter ochroma tin is co nde nsed chromatin and is t ranscriptionally inactive. In
electron micrographs, heterochromat in is e lectron de nse. An example of here-
rochrornan n is th e Barr bod y. which is found in female cells and represent s the in-
active X chromoso me.
2 . Euchromat in is dispersed chromati n and is tra nsc riptionally active.
3. A nucleosomc co nsists of DNA coiled arou nd htstones H 2A . H 2B. H J . and
H4. form ing an l l -nm -d ramerer chromati n fiber. A n uc leoso rne tha t has a
"beads-o n-a-s tr ing" appea ran ce is the basic unit of ch ro matin packagtng, These
l l -nm chnunartn fi bers ca n be packaged togethe r into 30-nm chro mat in fibers by
histone HI.
C . Ch romosomes contain some specialized nucleo tide seq uences.
1. Ce ntro mere s arc nucleotide sequences that mark the prim ary cons tr iction along
th e chromosome. Prote in co mplexes ca lled kinetocho res assemble at rhe cen-
tromere and bind mlcro rubules of the mitotic spindle durin g mitosis.
2 . Telc meres are nucleo tide seque nces (GGGlTA) located at the end of a chro mo-
some that allow replicat ion of DNA to its full length using the enzyme called
telcme rase,
3 . The replicat ion origin is a n uch..rond e sequence that serves as an originarion sire
of chromosome replicati on. Human chromosomes cont ain numerous replication
origins to ens ure rapid replica tion. In humans. DNA polyme rase a and & cata lyze
DNA replica tion. Orher DNA pol ymerases exist withi n the cell; namely DNA

1
2 Chapter 1

Table 1 ·1
Molecula r Transport Between N UC !l ' US and Cv rop lasm
Direction of Movement Mech anism

Ions Nucleus --) cytoplasm Passive trans port (diffusion)

Small molecules « 5000 d) Nucleus (-- cytoplasm No ATP hydrolysis
Proteins « 60,000 d)
mRNA Nucleus --) cytoplasm Active transport
IRNA Requires ATP hydrolysis
rRNA Requ ires bindi ng of RNA to protein s
with a signal sequence of 4-8
amino acids for recognit ion by the
nuclear pore com plex
Proteins [> 60 ,000 d) such as Nucleus (-- cyto plasm Active transport
nocrecprasmm . steroid Requires Al P hydrolysis
receptors. DNA and RNA Requires a signal sequence of 4-8
polymerases. gene regula- amino acid s for recogniti on by th e
tory protei ns , RNA-process- nuclear pore complex
ing proteins

Al P = adenosine triphosphate : mRNA ~ messenger RNA: rRNA - ribosomal RNA: tRNA = t ransfer RNA.

polym erase P and E:, which ca talyze DNA repair, and DNA po lvmeease v , whic h
catalyzes mitochond rial DNA replic at ion.
D. The n ucleo lus cons ists of portio ns of five pairs of ch romosomes [Le., 13, 14, 15, 21.
and ZZ ) that co nta in J;::en~ th at code for ribosomal RNA (rRN A) .ln h umans, RNA
polyme rase I catalyzes the formation of rRNA . Other RN A po lymerase, exist with in
the cell; namel y RNA polymerase II , wh ich catalyzes th e formation of messen ger
RN A (mRN A ), and RNA polyme rase Ill . which ca ta lyzes th e form ation of transfer
RN A (tRNA) . Ay elec tron nncroscopv, three rcgiom of th e nucleolus can be d ist in -
gu ished .
1 . The fibrill a r cente r is pale-stai n in g and co n ta ins transcription ally ina ctive DNA .
2. The densc fibrillar compone n t co ma ins rRN A in th e proc ess ofbeing svmhesued.
3 . T he gran ular componen t cont ains rRN A bo und to ribosomal protei ns beginni ng-
to mat ure int o ribos omes.

II. CYTOPLASM co nt a ins enzy mes for glvcolvsis, fatty ac id synthes is [i.e .• (an y aci d svn-
t h ase) , th ree reac t ions uf I he urea cycle (usin g anrininosucc inate syn thetase, argininosuc -
cmar c lyase, and arginase), glycogt"n synt h esis and degradati on , and protein syn th esis, as
we ll as inte rmed iates of meta bol ism and man y cofac ro rs ,

III . CYTOPLASM IC STRUCTURES

A. Ribosomes
1 . Ribosom es cons ist of of.QS (s mall) and 60S [larg e} subunits con ta in in g rRN A and
various proteins (Table ) -2).
2. They ate th e sites where transla tion of mRNA into an amin o add seq uence: [i.e.,
protei n svn t hcsis] occ urs.
3 . Rihoso mes 1Il,Iy clu ster alo n g a strand of mRNA to form a polyribosom e (or
 Cell Biology 3

Table 1-2
Rtbosornal Subc r ue
Number
Subunit rRNA Type of Proteins Functions

40S 18S Has binding sites for mRNA and tRNA

Binds to mRNA and finds the st art codon AUG
60S 55 , 5 .85, 285 Bind s to the 405 subunit after 405 subunit finds
th e sta rt codon AUG
Ha s peptidyl tr ansferase activity

rANA = ribosomal RNA; mRNA - messen ger RNA; tRNA '" traosfer RN....

po lysome] that is in volved in th e synthesis of cyto plasmic pro te ins (c .g., ac tin,
heruoglob ulin ].
I t I f( mn rER if rhe nascent pro-
4 , TIl ey lllay he di rected to the en dop lasmic ret iculum
te in co ntain s a hyd rophobic signal sequence at its amino terminal end. which is
c leaved in the rER IUIIlt'n by signal peptidase.
B. rER. This memhranous organelle contains ribosomes attached tc its cytoplas mic sur-
face by th e hind ing of ribophorin I and II to rbe ribosomal 60 S su bunit ,
1. It is th e site of synthesis of sec retory proteins [e.g., insu lin ), cell membrane pro-
tein s [e.g., recepto rs), and lyso somal er uvmes ,
2. It is the site of co-tra nslatio na l modification of prot cin s:
a. N elinkcd glvcosvlario n (add ilion of sugars to asparagine begin s in the rER an d
is completed in the ( Jo lgi com plex)
b. H yd roxylation of pro line and lysine d uring co llagen synt hesis
c. Cl eavage of the signal sequ ence
d. Fold ing of rhe nascent protein into three-dimensional co nfiguration
e . A ssoci ation of pro tei n sub un its into mulrimeric complex
C. Smooth endoplasmic retic ulu m (sER) is a membran ous organelle tha t contains no ri -
bosomes. It is invo lved in:
1 . Synthesis of membrane phosph olipid s {phosphandylcho hnc, sphing-omyelin ,
phosphandvlsenne. phosphat ldvlth anolatninc ], cholestero l, and ce ranude
2. Synthe sis of ste roid hormone s in testes. o vary, adrenal co rtex, and pla cen ta
3 . D ru g detoxification using cytochro me P HO' which is a family of h em e pro teins
(also called mixed-fu nct ion oxidase syste m ) that part icipates in h yd roxyla tion of
barbiturates, phen yro in , or benzopyren e (a carcinoge n found in c igaret te smoke ),
makes the m more soluble in water, and allows excretion into th e urine
a . Activati on of cytoc hrome P 450 hy one agent enhances the de tox ification of
ot he r agents . which has cl in ical implications.
b. In chronic alcoholics or newborns. large amounts of an esthesia age n ts arc
needed (whic h may he dangerous) beca use cvtocb rome P4'iO has been acti-
vated by detoxifying eithe r alcoho l o r break do wn prod ucts of feta l h ernoglob-
ulin , respect ive ly.
4. Fatty add elo ngat io n
5 . Calcium fluxes associated with muscle con tracuon
D. G olgi complexes are stacks of membran ous cisternae with a cis-face (co nvex) th at re-
4 Chapter 1

ceives vesicles of newly synthesized protein s from the rER and a rrens-face (co ncave)
that releases condensing vacuoles of posttranslationallv mod ified proteins.
1 . It is the site of pos ttranslaric nal modi fication of proteins , such as:
a . Comple tion of N vlinked glvcosv lat ion that began in the rER
b. O elinked glvcosvlatioru that is, ad d ition of sugars to serine by the enzyme gly-
cosvltransfcrase
c. Su lfa tion
d . P hosphorylation (p hosphorylation of man nosc funning mannose-e -phos-
phate occurs only in lysosomal enzymes)
2 . It is in vo lved in protein sorting and pac ka gin g.
a. Secre to ry protei ns (c.g., insulin ) are packaged into clarhrin-coared vesicles.
b. Cell membrane pro teins (e.g ., receptors) are packaged into nonclarhrin-
coated vesicles.
c . Lysosom al en zyme s are pac kaged into clarhr tn-coared ves icles after phospho-
rylation of mannose.
3 . It is involved in memb rane re cycling.
E. Mito chond ria
1 . Fun ct ion . Mitochondria are in volved in the produc tion of acetyl coenzyme A
(CoAl, the mcarboxvhc aci d cyc le, fatty acid ~-oxidation, amino HdJ oxidatio n,
and oxidat ive ph osph or ylation (wh ich ca uses the synthesis of adenos in e tri ph os-
ph ate (ATP ) d riven hy electro n transfer to oxyge n ].
8 . S ubstra tes are metabolized in the mitoc hond rial matrix to prod uce ace tyl
CoA, wh ich is ox idized by the tricarboxylic acid cycle to carbon d iox ide.
b . The energy released by this ox tdatton is cap tured by red uce d nicot inam ide
aden ine d inucl eot ide (NADH ) and flavi n adenine dmncleot ide (FA DH 1 ) .
NADlI and FADH 1 arc further oxidized, producing h ydrogen ion s and elec-
tron s.
c. 11,c electrons are transfe rred along the el ectron t ra nsport cha in , wh ich is ac-
companied by the outward pump ing of h ydrogen ions into the interme rnhrane
space {che m ios m ot ic t heory ).
d . The Fo subu nit of ATP synth ase forms a transmembrane hydrogen ion pore so
that hyd rogen ions ca n flow from the inre rmernbrane space into rhc ma trix,
where the F I subun it of AT P synthase catalyzesthe reac tion ADP + P; ~ AT P.
2 . C om ponen ts and con ten ts arc listed in Table 1-3.
3. Clin ical con sid eration s
8 . Led er 's h ereditary optic n europath y is ch ara ct erized hy progressive op tic
nerv e degeneration and is caused hy a m itochond rial DN A mutation in the
gene for su bunit 4 of th e NAD H dehydrogenase complex. Mitochond rial
diseases are mat erna lly inh erited and affect t issues that have a h igh requ ire-
me nt (Dr ATP (e.g., ne rve, muscle).
b . M yoclonic ep ileptic ra gged red fiber di sease is characterized by progressive
myoclonus (m uscle jerk in~), dement ia, an d hearing loss. Ir is caused by a mi -
tochond rial DNA mutation in the gene for t R N A for lysine.
c . Cyanide, ca rbo n mo no xid e, and antimyc in A inhibit the electr on tra nsport
chain and th us block AT P synt hes is.
d . O ligo mycin and vcn tu ricidin are antibio t ics that hind to AT P svn rhase and
thus block ATP synthesis.
F. Lysosomes are membrane- bound organe lles th at contain lysosoma l enzymes (a lso called
acid h ydrolase enzy mes) including cathepsin Band L (proreases), n uclease, 5' -nuclcotl-
 cell Biology 5

Tabl e 1-3
Compone n ts and Conten ts
Components Contents

Outer membrane Porin (a trans port prot ein that increases permeability to met abolic
sustrat es)
Int ermembrane s pace Hydrogen ions
Inner membrane Elect ron transport chain (NADH dehydrogenase, succi nate dehydrogenase ,
(folded int o cristae ) ubiQuinone-cytochro me c oxido reductase, cytochrome oxidase)
ATP synth ase (found on elementary particles)
ATP-ADP trensioca tor (moves ADP into the matrix and ATP out of th e
matrix)
Matrix compartment Tricarboxylic acid (TCA) cycle enzymes (except succi nate dehydrogenase )
Fatty acid j3-0xidat ion enzymes
Amino acid oxidat ion enzymes
Pyruvate dehydrogena se com plex
Carbamoylphosp hate synthet ase I
Ornit hine tra nscarbamoylase (part of urea cycle)
DNA, mRNA, tRNA, rRNA
Granules containing c alcium and magnesium ions

NAOH ~ reoocec nicotinamide adenine dinucleotide; m RNA - messenger RNA; rRNA "" ribosomal RNA; tRNA -
trans fer RNA; ATP = adenosine tri phos phate (ATP): AOP ~ adeno sine diphosphate (ADP).

dase, j3-g.llacIosiJoL'iC, fl-glUl,;u run i~, elvcostdase , aryl sulfatase, lipase , esterase, and acid
phos pha tase that func tion at pH 5. Mo:..t lysosomes funct ion in rracellu larly: however,
some cells (c.j{. , neutrophils. os rcocla-rs) release their lysosoma l cont ents extracellu larlv
1. Goi gi h ydrolase vesicle s buJ from the Golg i comple x and co ntain in active acid
hydrolase enzymes.
a . Golgi hyd rolase vesic les fuse with a late endosome. wh ich co n tains an 11 +-
AT Pase in its membrane that prod uct..-s a pH 5 en viron ment, which acti vates
th e acid hydrolascs.
b. A lat e endoso me may fu se with a phagocytic vacuo le formintl: a phagotvso..
some, which degrades material pha~ x:ytoseJ hy the cell .
c . A late endosome may fuse with an a utophagic vac uole forming an au"
toph agolv sosome, whic h degrades cel l organel le;
2. Resid ua l bod ies contain unchgesnble mater ial and may acc umulate with in a ce ll
as li pofus ci n pigmcn r.
3. C linica l cons iderations. The re are a n umber of gen et ic d iseases th at in vo lve rnu-
rat ion s of gvu cs for varlouslvsosomn l enzymes (acid h ydrola scs: Table 1·4 ).
G. Peroxisomes are membrane-bou nd organd ies.
1. Conten ts of peroxisom cs include:
a. Amino acid ox idase and hvdrc xvacid oxidase. wh ich produce h ydrogen per..
ox ide ( H I O I )
b. Ca talase and ot h er pe rox tdases tha t decom pose h ydrogen peroxide to warer
and oxyge n (H 20 2 --+ H 20 + 0 l )
c . Fatty acid fl-oxidati on en zymes rhat oxidiz e long-ch ain fatty acids (> 20 car-
bous] to short-cha in fany acids, wh ich arc transferred to mitoch ondria for
complete ox idat io n
2. C lin ical considerat ion. Adrenolcukcdvstroph v is a gene t ic d isease tha t in volves