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From www.bloodjournal.org by guest on June 15, 2017. For personal use only. Protein Synthesis in Human 1. Effect By T SEYMOUR protein Steroids WERTHAMER, HE IN VITRO taming medium phoid of Leukocytes and RNA and CAiu. CULTURE has been HICKS of rabbit demonstrated synthesis.1 and In liver Sterols AND cells in the however, both RNA suggesting and the protein target synthesis,2 while inhibiting the cell specificity of this hormone. of cortisol Inasmuch or cortisol as cortisol analogs, have been shown to is well known to be a “lympholytic that lymphopenia and cyte, that this seemed results at a specific cortin.5 This level latter synthesis of human regulator in the haps, may be binding its from inhibition feasible. the inhibition ultimately Furthermore, lymphocytes, the protein. of the result have with that its normal of of the therefore level lymphopenia.4 the possibility protein synthesis ratio the results MATERIALS plasma protein, does inhibit the in of the of plasma may act regulation levels the transprotein of as per- cortisol investigation to of the cortisol and related steroids and leukocytes and purified lympho- the AND plasma This undertaken cytes. presents increases result in agent,” lymphocytes. of action of of human mixed paper hormone of DNA,3 thus administration its binding if cortisol in vitro effect and mechanism sterols on protein synthesis This this synthesis In man, of lymphocytic number of a shift We in a cortisol coninhibition of lym- results in the destruction of the lymphocortisol is found to exist in the plasma and in equilibrium point would suggest control AMARAL LEONARD lymphoid to result cells, Lymphocytes: first phase of our investigation. METHODS MateriaLi Cortisol, Prednisolone and Testosterone respectively. 3-Methyl Cholanthrene Leo Soskind and J. was purchased chased from Julius from of All glassware human blood, used donated per respectively. International Chemical purchased Cholesterol ml. of Suspensions whole of for preparations by healthy blood ), and Nuclear Sigma, Sigma generously specific C14L-Ieuc1ne, Normal Corporation. Human and Vitarine, provided activity of Eagle’s TC submitted by 240 Drs. mC/mM, 199 was pur- allowed to sediment for and siliconized Hospital, a period Lymphocytes and sterilized. was heparinized of two hours the upper two thirds of the at 150 g. for 10 minutes. pipette-leaving just enough Whole (20 37 C. sedimented The superplasma to at accepted for publication May 15, 1969. Director of Laboratories, Methodist Hospital of Brooklyn; Clinical Assistant Professor, Department of Pathology, Downstate Medical Center, Brooklyn, N.Y. CARL HIcKs, M.S. : Research Assistant, Methodist Hospital of Brooklyn. LEONARD AMARAL, PH.D. : Director, Section of Research & Development, Department of Laboratories, Methodist Hospital of Brooklyn. SEYMOUR 348 WERTHAMER, February Leukocytes and incubations was employees of Methodist 5 ml. pipettes. After the sedimentation period, plasma was siphoned off, pooled, and centrifuged natant plasma was gently removed with a Pasteur in First from were Difco. Preparation units Carr, were and 6, 1969; M.D.: BLOOD, VOL. 34, No. 3 (SEPT.) 1969 From www.bloodjournal.org by guest on June 15, 2017. For personal use only. PROTEIN 349 SYNTHESIS cover the cap of leukocytes. The plasma was subsequently centrifuged at 1500 g. for 20 minutes. This step was sufficient to remove all the platelets from the plasma. For the cultures to contain leukocytes, the cap of leukocytes was genfly suspended in a volume of platelet free autologous plasma which would result in a cell number of 1 1 X 106 per ml. For pure lymphocyte cultures the cap of cells was gently suspended into a final volume of 3 ml. of platelet free plasma, and a pure suspension of lymphocytes, slightly contaminated by red blood cells was obtained according to the procedures described by Rabinowitz.#{176} The eluted lymphocyte suspensions were adjusted to 6.0 X 106 cells per ml. of plasma. The use of a single sample of 80 ml. of whole blood yielded enough cells for any one experiment. For those experiments involving the use of saline-washed cells resuspended in plasma or cells cultured After placed in centrifuged at with 2 ml. wash procedure 0.6 was 200 ml. cubated For ously dissolved 10-6, 10-s, are of or ml. 0.6 450 angle the use of the incubation of trypan detectable the for absence both cap of of used: cells were suspended and was then washed subsequently discarded. This the use of saline washed cells culture tubes to a final volume plasma, cultures described. the final volume subsequenfly All experiments and components and in saline -2 10 M. The of suspended were C of cent or performed 0.6 ml. and in- in duplicate earliest response the use at culture: 0.4 0.4 25 of the cells of two agents, ml. of 4 of C. the experimental final of TC 199 or 2 per agents of used of of ethanol), incubated period concentration culture cent were hour (previ- per -2 cultures ( contain- sterol concentration This the ml. hormone saline These and to 2.0 ml. a final suspension. 37, Period yield received lymphocyte 2 hours 3 ml. 14C-leucine), ethanol-to controls of Incubation of a final 3s per involving of the trypan above steps each at a incubation at 37 was C. For 10-s M. blue exclusion test for viability7 was routinely carried out after ( i.e., before column, after column, after centrifuging and after ). After each of the procedural steps, the viability, as judged by the criterion blue exclusion, remained upwards of 98 per cent, demonstrating that little injury had occurred. Used the were of Viability Procedures At the cells to be penicillin Determination The the g., Composition a period cultures each free The of leukocyte for represents those procedures the supernatant For the cultures involving plasma was added to the involving of Culture units and discarded, once. conditions following cultures. following 300 the at 200 platelet saline. the plasma, lymphocyte suspensions, 0.6 ml. of the suspension 2 ml. of saline was added. The culture tubes were supernatant cultures with Description The the repeated those under of centrifuged was triplicate ing g., in plasma, achieved or absence the adjusted tubes and of saline, resuspended of the obtaining into culture end of 0-4 C., After the the Incubation incubation of Cultures periods, ml. of 0.1 N NaOH g. for 10 minutes, placed resulting in the the in an supernatant ice bath and gently suspended in 2.0 ml. of 1 per cent acetic acid to disrupt the erythrocytes centrifuged at 200 g. for 10 minutes and the supernatant discarded. Three suesaline washes and centrifugations ( as before ) followed. The pellets of cells were 1.0 200 were pellets present,8 cessive in at cultures to resuspended centrifuged the cooled dissolving of the discarded, cells. These the were then neutralized with the equivalent amount of 0.1 N HCL, placed in an ice bath, the total protein precipitated with a final concentration of 5 per cent TCA, the precipitate subsequently collected and washed on Millipore filters ( 0.45 u pore size ) . These filters were then mounted on 2 inch planchettes and dried under infrared lights. The radioactivities of the above were obtained by the use of a Tracerlab 132 M Sealer gas flow counter (efficiency of 38 per cent ) . The data presented are the per cent alterations of the average total counts per minute per set in CPM of duplicate within or triplicate triplicate cultures cultures did to not that exceed of their controls. 4 per cent. In all cases the variations From www.bloodjournal.org by guest on June 15, 2017. For personal use only. 350 WERTHAMER, Fig. 1.-The effect of steroids and sterols on the incorporation of 14C leucine into human leukocyte total protein. An average of 6.5 X 106 leukocytes suspended in 0.6 obtained from hires of 9 different cell All incubations were formed molar agent p < in triplicate. concentration in the culture .001 for each experiments. formed lOco in no IP ft 1_I T I sicajIIllhIllUlIII iiii-I R E D N IS 0 L 0 N1 #{149}1R TDIt RUN METHYL ,3 donors. CH?LANTHREP4%.4 per- The final of each was series of I I -I ::cHoLEsTERo.L:.::::: per- I I I I I 40 30 20 10 0 4 C. resulted significant %INHIBITION inhibitions. CONTROL Fig. 2.-The effect of sterols and two concentrations of steroids on the incorporation of ‘4C leucine into lymphocyte total protein. Incubations _-4 are described in text. Results obtained at 10 NI and 10 M of steroid are presented as upper and lower bars, respec- ISOLO sTERoNE: I.’ tively. I :CHOLESTER9L: I I 30 20 per noted from different cell bations were triplicate. 0 series cent are obtained -l I The tions i-4-4 IHRAt 40 I I i I cul- Incubations at 25 and AMARAL CONTROL ml. of plasma were incubated at 37 C. under conditions described in the text. The per cent inhibitions noted are the averages HICKS, averages cultures 11 of donors. All men- for each performed p of inhibi- the < .001 in experiments. % INHIBITION RESULTS The effects cine were of the ( 3-methyl sterols ( cortisol, hormones cholanthrene, into leukocyte total protein observed when incubations are grade. Inasmuch as the leukocyte types, the inhibitions may above responses bers of the agents obtained described of each the cell leukocyte by in the the type thus population employed. To methods text. test ) results ) testosterone the incorporation and of the 14C-leu- . population be the leading may this on inhibitory at 37 C. ( Fig. 1 ) No inhibitions were performed at 25 and 4 degrees centriresult to the each possibility, of Rabinowitz,6 The prednisolone, cholesterol were composed of a summation possibility respond pure presented is in of various of that distinct a different suspensions mem- manner to of lymphocytes, incubated under in 2 indicate Figure cell unknown the conditions that the From www.bloodjournal.org by guest on June 15, 2017. For personal use only. PROTEIN 351 SYNTHESIS agents inhibited amino acid incorporation creasing the concentrations of the steroids cent inhibitions of amino acid incorporation. 2 indicate 3-methyl that the cholanthrene inhibitory are leukocyte incubations. This of leukocyte agents than population lymphocytes. lymphocyte, leukocyte The a pure procedures suspension tions for from glass of such to bead eluted and by and EDTA lymphocytes. ) results to incorporate the specific per the amino calculations affords into the type two non- it has of alteration for the determination in our study, the cell of these normal cent). elution acid and in the the separation of under the condi- However, that in a gross ‘4C suitable employed mathematical ( 90 neutrophil elsewhere columns some to the action portion of this Rabinowitz6 would afford which could be cultured presented no direct method is presently of neutrophils to the agents procedures is the total protein. Inincrease in the per of Figures 1 and to testosterone those produced that resistant largest leukocytes be cells suggests be more far, the population both ( data of the lymphocytes magnitude than comparison must By described by of neutrophils employed experience responses of a greater into lymphocyte resulted in no A comparison our been neutrophils in the protein. ability Although of the response of the following use deduction of such re- sponse. A prepared and B. The leukocyte total number determined for known for and the tures Triplicate equal cell preparation known and of of response was A. the suspension was of leukocytes control number neutrophils determined by the and were of triplicate equal cell of divided equally into and a cell differential experimental prepared. control number. the The leukocyte cells and experimental Incubations and leukocyte following cultures mathematical two parts, A analysis was to B were lymphocyte determination the agents culof the employed calculations: P = CPM Y = The number the per cent of lymphocytes of lymphocytes in the leukocyte culture which X leukocyte number divided which the per P X Y Z = by W \v =: - Z X = Performing ture the of lymphocyte equals the total total lymphocytes protein/lymphocyte. counts minute counts per minute of total leukocyte The counts per minute of protein for each these calculations their agents population. total of the corresponding experimental on the non-lymphocyte protein contributed ( mainly non-lymphocyte illustrate this Inasmuch teroP2 hibitions results protein of the leukocyte of that set. non-lymphocyte. and sets allows ( neutrophils lymphocyte ) of neutrophil) population. Table cul- one to deduce portion of the such experiments and calculations presented in Figure 3. A comparison of the Figure to Figure 2, suggests in reference to testosterone and 3-methyl cholanthrene, the inhibitions in the lymphocyte population are more marked than that observed leukocyte The of total equals by 100. present. The controls and effects of the of used cultures of part 1 serves to are that noted in the readily comparison. as cortisol,9 bind to plasma of amino acid testosterone,1#{176} proteins, the incorporation 3-methyl possibility produced cholanthrene1’ was investigated by these agents and choles- that the is dependent in- From www.bloodjournal.org by guest on June 15, 2017. For personal use only. 352 WERTHAMER, r r tu , D ( HICKS, AMARAL J I ISOL IIIICORT deduced efsteroids on incorporation of 14C leucine into neutrophil total protein. Methods involved in the fect the Fig. 3.-The of sterols and determination of presented ::.cHoLEsTERoL.::::I 40 30 20 10 the data are the averages obtained from the triplicate incubations different of cells individuals. from 3 0 % INHIBITION Table 1.-Comparison 14C Leucine Per into Cent of the Effects Total Protein Inhibition of Cortisol 14C of Agents Tested of Lymphocytes leucine Prednisolone on the Incorporation and Neutrophils Incorporation Produced 3-Methyl Testosterone of by: Cholanthrene Cholesterol Lymphocyte 21.4±1.4 14.6±1.7 17.8±1.9 17.0±2.5 22.0±4.3 Neutrophil 22 21 9 8 21 I CONTROL Fig. 4.-The effect of sterand steroids on the incorporation of 14C leucine into total protein by normal human lymphocytes cultured in the absence of plasma. Condi- lllc OR ols tions of culture in text. are from cells viduals. The are results I ISO llhJIIIIIIIJIIJIIJIJJIIllhIJJJltItIUhIIllllIllIllllllHIII L described presented METHYLCHOLANTHRENE the averages obtained triplicate cultures of from 3 different mdi- ::::cHoLEsTERoL I I I I I 40 30 20 10 0 % INHIBITION upon the the presence inhibitory acid into of plasma. conditions, of plasma. The of three actions the lymphocyte total protein The sterols, 3-methyl still inhibit amino acid results presented hormones are totally cholanthrene incorporation. on in Figure the 4 suggest incorporation of dependent upon and cholesterol, However, that amino the presence under these in the case of From www.bloodjournal.org by guest on June 15, 2017. For personal use only. PROTEIN 353 SYNTHESIS -II CONTROL Fig 5-The effect of ster 1111111 CORTISOL oids on the incorporation of 14C leucine into total protein IsoLE by human lymphocytes viously washed in saline TESTOSTERONEI cultured ing in medium autologous preand contain- i i 20 0 plasma. 40 30 0 % INHIBITION cholesterol, the magnitude in the presence To eliminate cells results in lymphocytes were subsequently cultured pear the to result of the loss in that the of in an increase as great as as those of of the the in the the above The the hormones, experiment results in Instead, of the of of the lymphocytes magnitude observed washing action plasma. hormones. that the to the containing washing actions not response saline in a medium inhibitory is to Figure 2). in these experiments that the washed 5 illustrate prevent inhibitions ( compare of plasma the possibility in saline in Figure of and presented saline saline inhibitions does not washes ap- of amino acid incorporation. Because the acid amino possibility was hypothesis of the the per cent suiting from tions in its cortisol Since these cortisol site are 38 per that of any two in their about and 3-methyl equal to 35 per and 3-methyl any agents ( 10 sterols, the different. the in no The concen- increase in significant increase rewhose final concentraM of lymphocyte in result ), site be increase to 2 of and may the tested ) over action. cholanthrene as only 3-methyl results in an cent since Figure of inhibition hormones observed previously cholanthrene action that ( see use the the of inhibition been a difference site of action or testosterone, 3-methyl of is has 6 suggest increases and tory indicates ( about expected site experiment those of between their which were Figure in this culture others either the in mediation different that hormones alone sented which is inhibition produced the simultaneous in that each by investigated tested tration l)y manner incorporation that The differs produced results from pre- all the cholanthrene, in conjunction with increase in the per cent inhibition. the sum of the for 3-methyl cent testosterone cholanthrene per cent ), respectively must be inhibitions cholanthrene different the from and inhibithat of hormones. DI5uSSI0N The results analog rene, and its lymphocyte presented prednisolone cholesterol, and leukocyte, in and inhibit this paper suggest testosterone, and the incorporation total protein. That that the the sterols, of amino these are hormones, cortisol, 3-methyl cholanthacid into human indeed inhibitions From www.bloodjournal.org by guest on June 15, 2017. For personal use only. 354 HICKS, WERTHAMER, I CONTROL lLc9.R AMARAL OLIIJIJIIJIIIIIII TIS IT E STOSTERON E 3 _________________ :#{149} c HO L E S I F RO L : ::::I Fig. 6.-The combinations ol ____________________ effect on corporation total protein IlllIIlHhlIlIIIllIIHIllIHIIllIllllIlllllhIII ___________________ of leucine by human phocytes. The centration culture of was description (iiIIIiiIILLLt%III1JIHIIIIIiII!iIiiJI1 is of sterthe in- final molar any 10. the of con- agent per Detailed experinent in the provided into lyni- text. . _ I I I 40 30 20 I I 10 0 % INHIBITION and not the caused by from 1 the ) the loss blue incubation 2 ) agents or and that precipitation of 1 per cent acetic loss of labelled studies protein have cell the cell injury, manner death or of handling, counts of agents hemagglutinin. ment with the lysis, is evident cultures employed serum total steroid dependence. precursor, for the cell washes following were neither of that lymphoid inhibits the the inhibitory effect Interestingly, cholesterol, cortisol1 cells. Furthermore, responses of possibility that produced in inhibit all sterols normally in- our the and in- indicating hydrocortison& prednisolone14 the lymphocyte as to phyto- are, in this respect, in agreeet al.’ found no dependence by cortisol studies, synthesis whose discarded yielded no activities these procedures. The results obtained in our study cited. However, whereas Makman those upon all acid supernatant protein during demonstrated synthesis hydrocortisone,15 gesting the cytolytic. Previous as from or tests indicate TCA-tungstate inhibit resulting employed exclusion periods eluding the no detectable well radioactivity the following: Trypan jurious of either we have both testosterone of lymphocyte nuclear structure observed and protein is not too a the sugdis- may have the same inhibitory effects. This suggestion appears plausible in that 3-methyl cholanthrene also has been noted in our studies to have such an effect ( although it apparently acts at a different site). Transcortin, a cortisol binding protein exists at a molar concentration of similar about and 10-i Doe M ) . in normal Their studies plasma have ( calculated shown that from this the protein data binds presented the by hormone Seal at From www.bloodjournal.org by guest on June 15, 2017. For personal use only. PROTEIN 355 SYNTHESIS equimolar concentrations. From the binding constant other studies,’6 the final concentration of cortisol binding is of the order of 10 M. At a concentration enhance of plasma, the inhibition value did not produce suggest, in the light incorporation by A similar binding site is not when in Our steroids with These suggest results means In therefore of a different conclusion, The molar of cortisol,17 synthesis. than that increased occur in of such result of the affinities to difcompete for if the would steroid twO of the above 3-methyl of complexes in the protein synthesis by present protein is normally would result of lymphopenia protein in- incorporation lymphocyte. condition of lymphocyte an observed in of plasma. of the binding protein of cortisol the by the when resulted inhibits lymphocyte since produced cholanthrene was also the absence site that effect, a difference action that such a complex, normally as a regulator of lymphocyte and inhibition that cholanthrene suggests may act of steroids, at a different inhibition suggest site never exceeded results, 3-methyl and complex system, testosterone. different testosterone a common amount of the cortisol therefore, administration formation as the that vitro protein vascular acid complex. However, present, of not of amino and with and 3-methyl cholanthrene acid incorporation in mechanism the cortisol-plasma in the human are suggest either inhibition. amino inhibition prednisolone and in the absence our results a transcortin-cortisol known. these cultures Unlike these conjunction the these at different sites, it would be expected results in no increase in its inhibitory results crease in the present our studies to inhibit by for presently both of action. per cent inhibition in more potent inhibitor. present that via apply were to inhibit in either steroid noted increase also data,16 mediated demonstrated to bind Whether or not cortisol sites protein complexes since an increase in their is by for the saturation M, cortisol did at 10 M. Since cortisol, of amino acid incorporation, transcortin may has been proteins.10 transcortin any of the lymphocytes mechanism Testosterone ferent plasma observed inhibition provided required of 10 higher in the would synthesis. SUMMARY The iii vitro effects steroids, cortisol, rate popultions It has been synthesis ably shown under mechanism as of inhibition indicating that these of the a protein cholesterol amino acid of sterols, all and 3-methyl incorporation. factor cholanthrene, However, is lower than a partial plasma port the hypothesis steroid hormones The physiologic to the condition that occurs used It hormones binding role of the of lymphopenia result has also requires in order cholanthrene in been the to the inhibition shown that presence achieve and synthesis in sepabeen investigated. of the of plasma, its effect. protein inhibitory presumThe sterol, in the absence of plasma, still inhibit in the case of cholesterol, the magnitude that observed dependence. the only 3-methyl testosterone on protein and leukocytes has agents conditions. steroid and cholesterol prednisolone and of human lymphocytes inhibition when the in the presence of The results presented of lymphocyte steroid is bound plasma protein-cortisol in man is discussed. plasma, perhaps therefore sup- protein synthesis by to a plasma protein. complex and its relation a From www.bloodjournal.org by guest on June 15, 2017. For personal use only. WERTHAMER, 356 SUMMARIO Esseva investigate e de le effectos steroides-cortisol, separate agentes in IN vitro de prednisolona, populationes usate resultava e 3-methyl-cholanthrena- testosterona-super le hormones rob steroide hypothese que occurre physiologic del solo complexo de lymphopenia in humanos We would Drs. Leonard like to thank Ornstein and e leucocytos del synthese le synthese de proteina in monstrate que omne le iste conditiones. Esseva etiam monstrate que le mechanismo inhibitori del hormones steroide require le presentia de plasma, presumitemente como un factor de ligation proteinic pro effectuar iste resultato. Le sterol-cholesterol e 3-methyl-cholanthrena-in le absentia de plasma inhibiva ancora le incorporation de amino-acido. Tamen, in le caso de cholesterol, le magnitude del inhibition esseva plus basse que illo observate in le presentia de plasma, possibilemente in consequentia de un partial dependentia ab le plasma. Le resultatos hic presentate supporta consequentemente de lymphocytos in le inhibition AMARAL INTERLINGUA steroles-cholesterol e HICKS, le inhibition quando de human. Esseva de proteina sub del synthese le steroide proteina lymphocytic es ligate a tin e cortisol e su plasmatic de proteina proteina de relation per plasma. con Le le condition es commentate. ACKNOWLEDGMENTS Miss Joanne Gallagher for her excellent Stanley L. Lee for their valuable advice. technical assistance, and REFERENCES 1. Makman, White, zation proteins Chem. M. Amaral, L., cytological cells of the 3. Amaral, renchyma of monal Seal, ities Steroids 6. S., sue and in aspects Doe, R. to of by tin: plasma. A in Aug. Book and Smith, ( ed. 3). Company, 12, 1967, the Transplantation Ono, T., blastoid Terayama, in human lymphoJ. and Isolation S. : by vivo. Incorporation normal Blood R., Sandberg, binding and of and leuke- 9: 1082, 1954. Schneider, Clin. H., vitro. Takoku, effects binding A. : Transcorprotein characterization. of of Seal, U. F., of hydro- phytohemagglutinin in 28: 1281, S., and Doe, 1. Isolation globulin. diethylstilbesterol Chem. the synthesis Endocr. 17. S., RNA on transin 1967. N. : Inhibitory upon Y. T. : Effect lymphocytes 5:317, NaKao, Effects : phytohemagglutinin- and human L. lymphocytes. 334. Oester, radioresistance of on p. S., and on J. Smith, drugs prednisolone induced phytohemagglutinin. W. corticosteroid IX J., I. Stefani, 15. 1968. A. C., and cytes. Biochem. Biophys. Acta 161:361, 1968. 16. DeMoor, P., and Heyns, W. : Cortisol binding affinity of plasma transcortin ( CBA) as studied by competitive adsorption. J. Slaumwhite, F. Forbes, 14. 23:811, 1967. hydrolases L-cysteine leukocytes Wissler, Blood 27:462, McGraw-Hill formation tis- Steven- cortisone Weissberger, radioactive 9. including A., R., Brittinger, G., HirschWeissman, C. : XII: redistri- Biol. 8. columns, and R. Handler, P., of Biochemistry anti-inflammatory induced lympho- J. Bio- p. 435. Lancet, of New testoster- Canad. Loebelt, Res. York, 13. Activ- Cancer. E., Cancer New and stimulated 37:412, Cell Pincus, of plasma. 1968. 12. White, A., E. L. : Principles N. leukocytes observations. K., and of acid bution cytes P. : In ) : Biological in Bresnick, 1964, hor- E. : Binding estradiol son, J. J., and Madix, J. C. : Distribution of radioactivity within hepatic cells after administration of labelled 3-methyl cholan- of Relation glass 1964. 7. Hirschhorn, horn, of Ann. Y. : Separation culture on 1966. B. 46:299, threne. cortisone leukocytes. T. (Eds. on of mouse liver paBiol. 35: 159, 1967. Some and 11. A 5:3526, Murphy, chem. parenchyma of polymorphonuclear monocytes one 1955. Rabinowitz, cytes, C. : Biochem. 10. in upon U. NaKao, effect S. : 59:907, C. and L. liver J. Cell A. influence of mouse DNA cells. Gordon, Y. Acad. Moriber, L. : The synthesis 5. and B., in vivo administration Biol. 35:3, 1967. J. Cell cortisone. mic and study following 4. Dvorkin, A. : Influence of cortisol on the utiliof precursors of nucleic acids and by lymphoid cells in vitro. J. Biol. 243:1485, 1968. 2. the H., 237:3136, human lympho- 1968. treated 1962. R. P.: from men. Cortisone plasma J. Biol. From www.bloodjournal.org by guest on June 15, 2017. For personal use only. 1969 34: 348-356 Protein Synthesis in Human Leukocytes and Lymphocytes: 1. Effect of Steroids and Sterols SEYMOUR WERTHAMER, CARL HICKS and LEONARD AMARAL Updated information and services can be found at: http://www.bloodjournal.org/content/34/3/348.full.html Articles on similar topics can be found in the following Blood collections Information about reproducing this article in parts or in its entirety may be found online at: http://www.bloodjournal.org/site/misc/rights.xhtml#repub_requests Information about ordering reprints may be found online at: http://www.bloodjournal.org/site/misc/rights.xhtml#reprints Information about subscriptions and ASH membership may be found online at: http://www.bloodjournal.org/site/subscriptions/index.xhtml Blood (print ISSN 0006-4971, online ISSN 1528-0020), is published weekly by the American Society of Hematology, 2021 L St, NW, Suite 900, Washington DC 20036. Copyright 2011 by The American Society of Hematology; all rights reserved.