'iote: This material is copyrighted. All rights reserved. Edward Goljan. :\1.0.

2002

~ Kava: sedative

.~ :\lelatonin: • sleep,. Rx of jet-lag

-.: St. John Wort: Rx of depression

:& Jojoba: cosmetic, hair growth

.,; Soy: phytoesrroger:

-&' Green tea: antioxidant. decrease cancer

~ Valerian: sleep

HIGH YIELD NOTES PHYSIOLOGY©

Proximal rena] tubule functions (see nephron diagram): • primary site for :\'a' reabsorpnon .• primary site for reclamation of HCO,-- loss of reclamation leads to type IJ proximal renal tubular acidosis due to loss of bicarbonate, • primary nephron site for synthesis of ammonia VIa enzymatic conversion of glutamine (non-toxic vehicle for carry l\H~ - In blood) into NH.;' + a-ketoglutarate .• primary site for reabsorption of glucose (cotransport with Na "), urea, amino acids. phosphate Functions of thin descending limb: only permeable to water. hence the urine becomes extremely hypertonic by the time it reaches the loop of Henle

Functions of thin ascending limb: Impermeable to water but permeable to Na and Cr. hence the COsm begins to decrease

Functions of thick ascending limb (TAL medullary segment): • generation of free water via the acn ve :\'a' -K - -2 Cl co-transport pump, • this pump is the most sensitive part of the nephron for damage due to tissue hypoxia, • blocked by loop diuretics, • pump also reabsorbs calcium (not PTH -enhanced)

Functions of the cortical TAL segment: • Na ~ /Cl' pump in early di stal tubule- Na and Ca (PTl-l-enhanced reabsorption) cations share the same channel for reabsorption, • blocked by thiazides

Functions of the macula densa: • interacts with the juxtaglomerular (JG) apparatus on the afrercn: arteriole .• increased Na 10 the urine inhibits renin release and vice versa

Functions of aldosterone-enhanced A TPase ~ a -'--K+ exchange pump in distal collecting tubule and collecting ducts: • Na - is reabsorbed in exchange for K-, • effect of increased distal delivery of '\-a ~ from more proximal acting diuretics (e.g .. loop diuretic or thiazide diureticj- * there is an augmented :\'a ~ ... K~ exchange, which may lead to hypokaiemia and increased reabsorpnon of bicarbonate (metabolic alkalosis)

Functions of the aldosterone enhanced HT/K+ ATPase pump and F ATPase pumps in the uintercalated cells in the collecting ducts (USMLE): primary sites for the excretion of excess H' :ons- dysfunction of the H- ... 'K- ATPase pump is the primary cause for type I distal RTA

:\ormal dilution of urine: • liOsm in the late distal collecting tubule/collecting duct IS normally --150 mOsmikg- primarily contains free water and a smaller amount of obligated water that must accompany solute .• when POsm is low. ADH is inhibited- absence of ADH causes the loss of free water in the urme, • positive free water clearance - * CH20 =0- V . COsmo where CH~O = free water clearance, V '" volume of urine in ml.zmin, eOsm =- obligated water. * to calculate COsm: COsm =COsm x V,'POsm, * a positive CH"O indicates dilution (free water is lost in the urine) ... example urine volume 10 ml., POsm 250 mOsm. lJOsm 150 mOsm: COsm = 150 x 10:250 = 6 mL CI!:O = : 0 ·6 = -4 rnl,

c"' :\ormal concentration of urine: • increase in POsm is a stimulus for ADH release. • ADH renders the late distal and collecting ducts permeable to free water (not Na. cannot reabsorb obligated wate;)- unne 1S concentrated .• negative CH:O (free water 1S reabsorbed back into the bloodl- e,g ..

. c

94

CORTEX

Proximal tubule

Filtrate

Isosmotic

HC03- regenerated

'\IEDLLL-\.

400 mOsm

Thin Ascending Limb

•

HPO .. ~-

::\'''U3 ... -=A=--H *

K~ or H~ HCO]- regenerated

ADH present (eoneen tra tion)

Collecting tubule

Functional Aspects oftbe ~epbron

A = Aldosterone

*H,-O =- free 'water: mostly generated in Xa ~1K~/2Cr cotransport pump ADH = antidiuretic hormone

TAL = thick. ascending limb (diluting segment) OCT = distal convoluted tubule

H'" = hydrogen ion that can be secreted with HP04- (H:P04), which is titratable acidity or :"."H, ('"aO)

Hypertonic

Loop of Henle

1200 mOsm

ADH absent (dilution) positive CH20

Proximal tubule: • carbonic anhydrase inhibitor- blocks reclamat.on of bicarbonate. which lS excreted as '\aHCO, or KHeQ,: +proximal renal tubular acidosis, + loss of K. and '\a'

Ascending tubule: • loop diuretic- blocks :\2.~,X-,,'2C1" cotransport pump: +~r.1pai",s generation of free water. + (:8.:;::: :"::1'_ :5 a.so .ost ir: the urine (Rx of hypercalcemia), + hyponatremia, hypokalemia, metabolic alkalosis. • thiazide diuretic- blocks ): 8. ~'C;- pump: + allows calcium reabsorption wit;' the help of parathorrnone (useru. .r: caic:u:n St.O:1C formers). * hyponatremia. hypokalemia, metabolic alkalosis

Distal/collecting tubule: • spironolactone- blocks aldosterone (A) purr::ps: + blocks :\ a K - exchange pt.:.mp. + O:OCK~ :--l~ K - ATPase exchange pump, + K~ sparer. + normal anion gap metabolic acidosis

:'\ote: This material is copyrighted. All rights reserved. Edward Goljan. 'LD. 2002

urme volume 10 ml., POsm 300 mOsm'kg. UOsm 900 mOsm'kg: COsm = 900 x 10,'300 = 30 ml., CH,O = 10 . ]0 = -20 mL. • ability to concentrate urine is the first abnormality in renal failure

_i/ Central and nephrogenic Dl: • both have low UOsrn and increased POsm .• central DI snows> 50% increase m l:Osm with administration of vasopressin .• nephrogenic DI shows <50°'0 Increase In COsm with vasopressin

,~ Respiration changes with increasing altitude: • respiratory alkalosis. hypoxemia due to decreased atmospheric pressure not a decrease in % oxygen in air. • increase in 2.3 BPG right sbfb the ODe

~ Effect of VIP (vasointestinal peptide) and enkephalins on Gl tract: • VIP- relaxes smooth muscle, increases intestinal secretion. increases pancreatic secretion .• enkephaiins (opiatesjcontract smooth muscle. decrease intestinal secretion

~ Know peol dissociation curve

~ Inhibin: • synthesized In Sertoli cells m seminiferous tubules .• negative feedback with FSH, • increased If seminiferous tubules are destroyed, • normal If Leydig cells arc destroyed. since testosterone has a negative feedback with LH

,",' Calculations:. alveolar ventilation- PA02 = PiOz - PACO:,IR. where PiOc equals % oxygen x 713) and R is the respiratory quotient that normally equals 0.8. subtract PaOe from PAO=, and y'OU have the A-a gradIent. F1Ck's equation for cardiac output- cardiac output ml.rrnin e- oxygen consumption + oxygen in pulmonary vein - oxygen in pulmonary artery'

:._' Hormone increasing/decreasing gastric secretion: • gastnn .• hormones inhibiting acid secretion- secretin and gastric Inhibitory peptide

.~ Placental anatomy/physiology: • maternal surface has slightly bulging areas called cotyledons. which are covered by a layer of decidua basalis •• fetal surface is entirely covered by the chorionic plate- chorionic vessels converge with the umbilical cord, which is composed of 2 umbilical arteries (venous blood returning from the fetal heart) and 1 umbilical vein (carries oxygenated blood from the placenta), • chorionic villus/umbilical cord- * chorionic villi project I:": the mtervillous space. which contains maternal blood from which oxygen is extracted: spiral arteries from the uterus empty into the space. * chorionic villi are lined by trophoblastic tISSUe: outside layer IS composed of syncytiotrophoblast: synthesizes heG and human placental lactogen (growth hormone of pregnancy) and inside layer is composed of cytotrophoblast: clear cells. the intenor of the chorionic villus has fetal blood vessels, which coalesce to form the chorionic vessels that converge with the umbilical cord .• the umbilical cord contains 2 umbilical arteries (contains deoxygenated blood exiting the fetal heart and returning to the placenta) and 1 umbilical win (contains oxygenated blood)

., Effect of increased total peripheral resistance on vascular function curve: • same as the venous return curve, which depicts the relationship between venous return and nght atrial pressure .• increasing TPR (vasoconstriction) decreases venous return to heart. decrease nght atnal pressure. and decrease cardiac output- this causes a counterclockwise rotation of the vascular function curve .• decreasing TPR (vasodilation) causes a clockwise rotation of the curve- venous return increases. right atrial pressure increases, and cardiac output increases

-if hCG:. has luteinizing hormone activity, .• keeps corpus luteurn of pregnancy synthesizing progesterone until 8-10 wks and then placenta takes over that function

~ Stimulation of histamine H2 receptors: increases secretion of acid by parietal cells

.. ~ GI peptide injected into cerebrospinal fluid that increases appetite for carbohydrate: somatostatin- this normally is made in hypothalamus where it functions to inhibit growth hormone. hence inhibiting GH causes hypoglycemia and the body would crave more carbohydrate to increase gl ucosc lewis

- Atrial natriuretic peptide: mediated by guanylate cyclase. increased If ieft or right a::-1;.:;-1' is VOlume overloaded

95

:"':ote: This material is copyrighted. All rights reserved. Edward Goljan. :\-t.D. 2002

,/:"

Central diabetes insipidus: injecting vasopressin causes an increase In urme concentration (POsm) and decrease in urine volume

Location of angiotensin converting enzyme: • pulmonary capillaries .• increased In sarcoidosis .• non-competitive inhibition by' ACE inhibitors

Vitamin D type in kidneys: • 1.25 (OH)2D" • kidney has 1 a-hydroxylase in proximal tubules (PTH stimulates synthesis) .• renal disease )vICe of hypovitaminosis D

Effect of thyroid hormone excess on bone: osteoporosis- increases bone turnover resulting m loss of bone mass

Primary site for temperature regulation: • anterior hypothalamus, • fever is due to release of pyrogens stimulating LL-l release from macrophages- * IL-I increases synthesis of PG E~ tn the anterior hypothalamus, *PGE2 raises the hypothalamic set-point (normal core temperature viewed as too 10\\). hence anterior hypothalamic reactions of heat generation prevail. • heat generatrng mechanisms if core temperature is below set-point include- -increasing the release of thyroid hormones (increases metabolic rate), =vasoconsrriction of skm vessels (sympathetic stimulation 0,' ex-receptors in smooth muscle), -syrnpathetic stimulation of l3-receptors in brown fat (increases metabolic rare and heat production), =shivering (most effective system, center located in posterior hypothalamus, leads to activation of a and y motoncurons innervating skeletal muscle) .• heat dissipating mechanisms (coordinated in posterior hypothalamus) If core temperature is above setpoint mclude- -reducing sympathetic tone (vasodilatation) of skin vessels leading to shuntmg of blood through venous plexus in the skin, +increased sympathetic activity of cholmergic fibers innervating sweat glands leading to sweating

Cause of decrease in jugular venous pulse on inspiration: • decrease in lung compliance (c.g .. sarcoidosis), • decreased volume of air in the alveoli leads to less compression of vessels in the interstitial tissue. hence they fill up better and dram blood off easier

Iron reabsorption: • meat has heme iron, which is ferrous and easy to reabsorb in the small intestine (primarily the duodenum): * once absorbed into enterocytes, heme is cnzyrnaticail .... degraded to release iron, * most of the iron is diverted to storage as ferritin in the enterocyte (called apoferritin), while a small amount is delivered to plasma transferrin, the circulating binding protein of iron .• plants have non-heme iron, which is in the ferric state: * non-heme iron first binds to mucin in the stomach (renders it soluble), * the mucin-nan-heme complex then binds to proteins on the mucosal surface of the duodenum for transport into the cytosol where it IS enzymatically degraded to release iron, * a cytosolic protein called mobilferrin transfers the non to the mucosal rerritm stores or to transferrin in the plasma, • when body stores of iron are replete. mucosal cells an: shed With their stored mucosal ferritin in order to prevent Iron overload and transfemn synthesis in the liver is decreased .• when body stores are depleted, most of the i:-01"': IS directed [0\\·a:-05 transferrin in the plasma and transferrin synthesis in the liver is increased.

Know ventricular volume curve and where different heart sounds are located: c.g., Sc (closure of A\' and PV)

Renal reabsorption curve: glucose 1S most often asked about

Destroy hypothalamus. what hormone increases: prolactin- no dopamine to inhibit Effect of carotid massage: slows hean rate

Major site of water reabsorption in the GI tract: in descending order: jejunum, Ileum, colon Most effective nephron site for acid excretion: proximal tubule of the kidney

:\"ephron site for ADH effect: collecting tubule

:\" ephron site for greatest generation of free water/most susceptible to ischemia: thick ascending limb In the medulla

Constriction of the efferent arteriole: increases the GFR and decreases renal plasma 110\\ Filtration fraction in the glomerulus: FF = GFR./RPf; constrict afferent arteriole, no change ;;-: FF (Y GFRv RPf). constrict efferent arteriole increases FF (':' GFR' ..... RPF). increase plasma protem

,i'"

96

:-':ote: This material is copyrighted. All rights reserved. Edward Goljan. ,!\LD. 2002

concentration. hence increasing 1l:GC. decreases the FF (.,j_ Gf'R'no change RPF); decreasing plasma protein concentration. hence decreasing TCGC, increases the FF (' GFR'no change RPF); constricting the ureter. decreases PBS. hence decreasing GFR and the FF without affectmg RPF ('" GFR'no change RPF)

.~ Starling equation for GFR mllmin: GFR = Kf [(PGC - PBS) - 11GC]' where Kf = filtration coefficient In ml.rnin . nun Hg, PGC = hydrostatic pressure in the glomerular capillary in mm Hg (pushes fluid Q;Jt of the capillary; e.g .. ~45 mm Hg; it is increased if the efferent arteriole IS constricted and decreased if the afferent arteriole IS constricted), P8S = hydrostatic pressure in Bowman's space in mrn Hg (pushes fluid into the capillary; e.g .. -10 mm Hg), and 11GC = the oncotic pressure in the glomerular capillary in rum Hg (brings fluid back into the capillary: e.g .• -19 rum Hg): the net filtration is the algebraic sum of the above 3 parameters (in the above example. the net pressure would be -10 - 45 -19 = ... 16 rnm Hg)--fluid should move out of the glomerular capillancs: question: what Starling force changes \0 produce a net ultrafiltration of zero (answer: 71:(;(. which becomes increased [pulls fluid back into the glomerular capillary: using the above example-s- 10 - 45 - 35 = 0 net pressure]; it 1S the only factor that does not remain constant along the length of the capillary owing to differences in protein concentration: the high 7I:GC at the end of the glomerular capillary extends into the pcritubular capillaries that surrounds the nephrons and allows for the reabsorption of solutes).

:it Graph of pH, PC02, bicarbonate: PCOz 40 rom Hg is normal, bicarbonate 24 mliq/I. is normal

PC02 isobars 60

Bicarbonate

24 mEq/L

7.0

7.40

7.8

j;!'

Arterial pH

• patient A- acute respiratory acidosis with very little bicarbonate compensation .• patient Bchronic respiratory acidosis where there is more compensation by increasing bicarbonate (rnetabohc alkalosis) and pH comes closer to normal range .• patient C- metabolic alkalosis where there 15 compensation by increasing PCO~ (respiratory acidosis) • patient D- metabolic acidosis where there is compensation by decreasing PCO:! (respiratory alkalosis). • patient E- acute respiratory alkalosis where there is very little drop in bicarbonate for compensation (metabolic acidosis). • patient F~ chronic respiratory alkalosis where there 1S good compensation by dropping bicarbonate (metabohc acidosis) and pH is closer to normal

:\egative charge of GB:\L heparan sul fate

S, heart sound correlates with C wave in the jugular venous pulses (C wave is closure of the tricuspid valve): know the jugular venous pulses: see box in cardiovascular chapter on physical CWgnOS1S

Carotid massage: • decreases heart rate and increases vasodilatation (carotid sinus baroreceptor innervated by the LX and Xth nerve .• impulses generated III this receptor inhibit tonic discharge of

97

Xote: This material is copyrighted. All rights reserved. Edward Goljan. :vI.D. 2002

the vasoconstrictor nerves and excite vagal innervation of the heart producing vasodilatation, venodilaticn. drop in blood pressure and heart rate, and a decrease In cardiac output)

.i: 3% Hypertonic saline and effect on POsm and ADH levels: • 3% hypertonic saline mcreases POsm, • mcreased POsm stimulates the release of ADH (increases) .• atrial natriuretic peptide is also stimulated and normally does inhibit ADH release. however, hypertonicity overrides A?\"P

~ Patient walking briskly on a hot day: no increase In rectal temperature, vasodilatation 0:- vessels in skin

._. Marathon runner on a hot day: • increase in rectal temperature .• vasodilatation of vessels III skin

.~ Vagus nerve functions in the stomach: • increases gastric acid secretion and gastnc motrl iryvagotomy' decreases acid secretion and motility, • the 'Vagus nerve represents the parasympathetic innervation of the stomach and duodenum

';,' Sympathetic nervous system in the stomachJduodenum: • inhibits sccrenon and motility, • Sympathetic fibers are located in the celiac plexus from T5- T9

:.it Motilin: hormone modulating GI muscle activity

:.it 0.9% normal saline with Kef is the crystalloid solution that is most often used to replace gastric loses

:g Creatinine clearance (CCr): • creatinine clearance fonnula- * CCr = COsm x V··' POsm. where V ..= volume of a 14 hr urine collection 10 ml.zrnin, • creatinine 1S not a perfect clearance substance .• causes of a decreased CCr-" increasing age, * inadequate 14 hr urine collection: decreases urine volume in the numerator, " renal failure .• causes of an increased CCr- ,. normal pregnancy: increase m plasma volume increases GFR and eeL * early diabetic nephropathy:

. .; FE-:\" a ~ (fractional excretion of sodium): • useful in the workup of oliguria- FE?\a ~ = (L\' a' x PCr) i (P'Na - x CCr) x 100 .• values < 1 indicate intact tubular function .• values> lrusually :> 1) indicate tubular dysfunction

':,0 :\lyocardial physiology: • cardiac hypertrophy increases wall stress. sinus tachycardia reduces fi:ling of the coronary arteries, • increasing heart rate raises myocardial oxygen COnSU:ilp::oncardiac muscle uses i)-oxidation of fatty acids for energy .• \'enoconstnctlOn- increases preload alone, • restrict salt and water intake-- decreases preload alone .• increase cardiac contracuhry- no change m preload/afterload

,~. ~ormal gas physiology at the tissue level: • CO: derived from tissue enters the RBC and

combines with HeO via carbonic anhydrase to form H2C03, • H~C03 dissociates into H"" and HCO~ the latter leaving the RBC in exchange for CI- anions, • H~ combines with oxygenated Hgb (HgbOJ, which releases 0: <Bohr effect), • 02 leaves the RBC dissolves In plasma and increases

capillary PO:

- Xormal gas physiology at the pulmonary level: • Alveolar 0:, diffuses mto the plasma of the pulmonary capillary owing to a greater partial pressure of alveolar O:~, • 0: enters the RBC and

combines with ferrous ions on deoxyhemoglobin to form oxyHgb and H-, • RCO:" enters the RBC from the plasma and combines with R+ to form HFO" • Cl" anions leave the RBC to counterbalance the entry of HCOj', • H2COj dissociates into CO~ and H:O: CO2 leaves the RBC and dissolvc« in the plasma to increase the PCO> which enters the alveoli for excretion

:;.,. Important anatomical considerations for urine control: • urogenital diaphragm- " muscles 0:' the urogenital d:aphragm are the deep trans .... ·erse perineal and sphincter urethra muscle. ~ location of the urogenital diaphragm on a cystourethrogram is just distal to the prostate m a male. " 10ca::0:1 of the urogenital diaphragm on a cystourethrogram in a female is the beginning of urethra where i: ex.ts the bladder, • functions of the detrusor muscle- " relaxed: storage of urme lTI the bladder. " contracted: emptying of bladder. • functions of the sympathetic in bladder control- " relaxes the detrusor muscle: aids in urine storage In the bladder. * contracts internal sphincter: increases urine

98

Xote: This material is copyrighted. All rights reserved. Edward Goljan. :M.D. 2002

storage .• functions of the parasympathetic system- '" contracts the detrusor muscle: empties the bladder. * relaxes the internal sphincter muscle by blocking sympathetic inhibition: 2.110\\<; emptying of the bladder

.~ Thyroid hormone: • diffuses into cytosol. • binds to receptors in the nucleus: DXA binding domain has zinc fingers, other hormones with similar binding: steroids. retinoic acid, vitarnm D

"" Angiotensinogen: • synthesized in liver. • increased synthesis from estrogen

~ Action potential curve of cardiac muscle: know what part corresponds to the T wave (right at the end)

H Ventricular and aortic pressure curve: where is S2 (beginning of diastole)

c' Substance requiring ATP for reabsorption in the small bowel: • glucose, • ATP required for cotransport with Na

~ :\lajor vehicle for carrying CO2 in blood: bicarbonate (-70%)

"i: Volumes/capacities not directly measured by spirometry: • residual volume (RV): volume of air left over after maximal expiration, • total lung capacity (TLC): amount of air in a fully expanded lung, measured with a nitrogen or helium dilution method, • functional residual capacity (FRe): total amount of air in the lungs at the end of a normal expiration (end of the tidal volume lTV]), obtained by a helium dilution technique or body plethysmography

it Forced vital capacity (FVC), forced expiratory volume in 1 second (FEVh~<) and FEV1 se cfFVC: • FVC is the total amount of air expelled after a maximal inspiration: the RV IS what 1S left over at the end of maximal expiration: normal FVC is 5 liters. • forced expiratory volume in 1 second (FEV1,ec): FEV]>c" is hov .... much air a person can expel from the lungs in 1 second after a maximal Inspiration, normal FEV1sec IS 4 liters. • ratio of FEVh~ciFVC is normally 0.80: 4 liters 5 liters = O_SO, • peak expiratory flow meter: outpatient method of evaluating forced vital capacity (FVC). commonly used by asthmatics to evaluate their airv ... ays

1: Method of measuring RV: • subtract the expiratory reserve volume (ERV) from the functional residual capacity (FRC), • ERV is the amount of air forcibly expelled at the end of a norma. expiration {end of the TV),. FRC = ERV +- RV, therefore. RV = FRC - ERV

~ Diffusion capacity (DLeo): • DLeo is the method of measunng the ability of a gas to diffuse through the alveolar/capillary interface: utilizes carbon monoxide (CO) .• DLeo 1S primarily dependent on the following parameters: CO reaching the alveoli (decreased due to VQ mismatches: e.g.. atelectasis, COPD), CO crossing the alveolar/capillary interface (decreased due to pulmonary fibrosis or fluids in the interface)

99

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-~-, Seccnas

_.. 3 L

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4. -\'\

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v- __ -_

B C

• TL eve -",:__/_L_L__Ij'-+---f-L__jj'---.L-J'---J

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FIGCR£ 11- L Schematic of the normal lung volumes and capacities and the forced expiratory volume 1 second (FEV:,...J and forced vital capacity (FVq in a normal person CA.), a person with restrictive lung disease (B), and a person with obstructive lung disease CC)- FRC (functional residual capacity) represents the volume of gas ~hat remains in the lung at rest at the end of a normal respiration and is the most reproducible part of a pulrnonary function test. since it does not require patient effort. Person A (normal individual) has an ITV,,,,, of 4 L and all PvC of 5 L with F£\",ec'F'v'C ratio ~ 0.80. Person B with restrictive :ung disease has a "miniaturized" curve of person A ...... ote that the FE\--":o"", and F\/C are the same (3 L) owing to the increase in elasticity in the lungs. hence FEV"""iFVC ratio = :.0. Person C with the obstructive pattern is having difficulty in expelling air from the lungs owing to decreased lung elasticity. FEV;soc '" : L. F\'C = 3 L, and FE\i:,,~c.'FVC ratio = 0.33. Both people B anc C have reduced values for FEV",."" and F'vT when compared with the normal person: however. person B with restrictive .ung disease has values oetwecn those of tne normal .ndividuai and person C with obstruction, Person B has decreased compiiancc :n the .ungs, hence jess air enters the lungs. but OWing to the increased elasticity. the air that is in the lungs :s expelled faster. Person C ;--,as increased compliance (air easily enters the lungs): however. reduced eiasticity causes the lungs to trap air on expiration. hence FEY",,", and F'v'C are much lower than if'. person B anc the r::::V'''CifVC ratio :5 reduced, TLC. total lung capacity: 7\:. t.cai volume: vC. vital capacity.