a rXiv:h ep-ph/9212266v116Dec1992SLAC–PUB–6017WIS–92/99/Dec–PH December 1992T/E The Subleading Isgur-Wise Form Factor χ3(v ·v ′)to Order αs in QCD Sum Rules Matthias Neubert Stanford Linear Accelerator Center Stanford University,Stanford,California 94309Zoltan Ligeti and Yosef Nir Weizmann Institute of Science Physics Department,Rehovot 76100,Israel We calculate the contributions arising at order αs in the QCD sum rule for the spin-symmetry violating universal function χ3(v ·v ′),which appears at order 1/m Q in the heavy quark expansion of meson form factors.In particular,we derive the two-loop perturbative contribution to the sum rule.Over the kinematic range accessible in B →D (∗)ℓνdecays,we find that χ3(v ·v ′)does not exceed the level of ∼1%,indicating that power corrections induced by the chromo-magnetic operator in the heavy quark expansion are small.(submitted to Physical Review D)I.INTRODUCTIONIn the heavy quark effective theory(HQET),the hadronic matrix elements describing the semileptonic decays M(v)→M′(v′)ℓν,where M and M′are pseudoscalar or vector mesons containing a heavy quark,can be systematically expanded in inverse powers of the heavy quark masses[1–5].The coefficients in this expansion are m Q-independent,universal functions of the kinematic variable y=v·v′.These so-called Isgur-Wise form factors characterize the properties of the cloud of light quarks and gluons surrounding the heavy quarks,which act as static color sources.At leading order,a single functionξ(y)suffices to parameterize all matrix elements[6].This is expressed in the compact trace formula[5,7] M′(v′)|J(0)|M(v) =−ξ(y)tr{(2)m M P+ −γ5;pseudoscalar meson/ǫ;vector mesonis a spin wave function that describes correctly the transformation properties(under boosts and heavy quark spin rotations)of the meson states in the effective theory.P+=1g s2m Q O mag,O mag=M′(v′)ΓP+iσαβM(v) .(4)The mass parameter¯Λsets the canonical scale for power corrections in HQET.In the m Q→∞limit,it measures thefinite mass difference between a heavy meson and the heavy quark that it contains[11].By factoring out this parameter,χαβ(v,v′)becomes dimensionless.The most general decomposition of this form factor involves two real,scalar functionsχ2(y)andχ3(y)defined by[10]χαβ(v,v′)=(v′αγβ−v′βγα)χ2(y)−2iσαβχ3(y).(5)Irrespective of the structure of the current J ,the form factor χ3(y )appears always in the following combination with ξ(y ):ξ(y )+2Z ¯Λ d M m Q ′ χ3(y ),(6)where d P =3for a pseudoscalar and d V =−1for a vector meson.It thus effectively renormalizes the leading Isgur-Wise function,preserving its normalization at y =1since χ3(1)=0according to Luke’s theorem [10].Eq.(6)shows that knowledge of χ3(y )is needed if one wants to relate processes which are connected by the spin symmetry,such as B →D ℓνand B →D ∗ℓν.Being hadronic form factors,the universal functions in HQET can only be investigated using nonperturbative methods.QCD sum rules have become very popular for this purpose.They have been reformulated in the context of the effective theory and have been applied to the study of meson decay constants and the Isgur-Wise functions both in leading and next-to-leading order in the 1/m Q expansion [12–21].In particular,it has been shown that very simple predictions for the spin-symmetry violating form factors are obtained when terms of order αs are neglected,namely [17]χ2(y )=0,χ3(y )∝ ¯q g s σαβG αβq [1−ξ(y )].(7)In this approach χ3(y )is proportional to the mixed quark-gluon condensate,and it was estimated that χ3(y )∼1%for large recoil (y ∼1.5).In a recent work we have refined the prediction for χ2(y )by including contributions of order αs in the sum rule analysis [20].We found that these are as important as the contribution of the mixed condensate in (7).It is,therefore,worthwhile to include such effects also in the analysis of χ3(y ).This is the purpose of this article.II.DERIV ATION OF THE SUM RULEThe QCD sum rule analysis of the functions χ2(y )and χ3(y )is very similar.We shall,therefore,only briefly sketch the general procedure and refer for details to Refs.[17,20].Our starting point is the correlatord x d x ′d ze i (k ′·x ′−k ·x ) 0|T[¯q ΓM ′P ′+ΓP +iσαβP +ΓM+Ξ3(ω,ω′,y )tr 2σαβ2(1+/v ′),and we omit the velocity labels in h and h ′for simplicity.The heavy-light currents interpolate pseudoscalar or vector mesons,depending on the choice ΓM =−γ5or ΓM =γµ−v µ,respectively.The external momenta k and k ′in (8)are the “residual”off-shell momenta of the heavy quarks.Due to the phase redefinition of the effective heavy quark fields in HQET,they are related to the total momenta P and P ′by k =P −m Q v and k ′=P ′−m Q ′v ′[3].The coefficient functions Ξi are analytic in ω=2v ·k and ω′=2v ′·k ′,with discontinuities for positive values of these variables.They can be saturated by intermediate states which couple to the heavy-light currents.In particular,there is a double-pole contribution from the ground-state mesons M and M ′.To leading order in the 1/m Q expansion the pole position is at ω=ω′=2¯Λ.In the case of Ξ2,the residue of the pole is proportional to the universal function χ2(y ).For Ξ3the situation is more complicated,however,since insertions of the chromo-magnetic operator not only renormalize the leading Isgur-Wise function,but also the coupling of the heavy mesons to the interpolating heavy-light currents (i.e.,the meson decay constants)and the physical meson masses,which define the position of the pole.1The correct expression for the pole contribution to Ξ3is [17]Ξpole 3(ω,ω′,y )=F 2(ω−2¯Λ+iǫ) .(9)Here F is the analog of the meson decay constant in the effective theory (F ∼f M√m QδΛ2+... , 0|j (0)|M (v ) =iF2G 2tr 2σαβΓP +σαβM (v ) ,where the ellipses represent spin-symmetry conserving or higher order power corrections,and j =¯q Γh (v ).In terms of the vector–pseudoscalar mass splitting,the parameter δΛ2isgiven by m 2V −m 2P =−8¯ΛδΛ2.For not too small,negative values of ωand ω′,the coefficient function Ξ3can be approx-imated as a perturbative series in αs ,supplemented by the leading power corrections in 1/ωand 1/ω′,which are proportional to vacuum expectation values of local quark-gluon opera-tors,the so-called condensates [22].This is how nonperturbative corrections are incorporated in this approach.The idea of QCD sum rules is to match this theoretical representation of Ξ3to the phenomenological pole contribution given in (9).To this end,one first writes the theoretical expression in terms of a double dispersion integral,Ξth 3(ω,ω′,y )= d νd ν′ρth 3(ν,ν′,y )1Thereare no such additional terms for Ξ2because of the peculiar trace structure associated with this coefficient function.possible subtraction terms.Because of theflavor symmetry it is natural to set the Borel parameters associated withωandω′equal:τ=τ′=2T.One then introduces new variables ω±=12T ξ(y) F2e−2¯Λ/T=ω0dω+e−ω+/T ρth3(ω+,y)≡K(T,ω0,y).(12)The effective spectral density ρth3arises after integration of the double spectral density over ω−.Note that for each contribution to it the dependence onω+is known on dimensionalgrounds.It thus suffices to calculate directly the Borel transform of the individual con-tributions toΞth3,corresponding to the limitω0→∞in(12).Theω0-dependence can be recovered at the end of the calculation.When terms of orderαs are neglected,contributions to the sum rule forΞ3can only be proportional to condensates involving the gluonfield,since there is no way to contract the gluon contained in O mag.The leading power correction of this type is represented by the diagram shown in Fig.1(d).It is proportional to the mixed quark-gluon condensate and,as shown in Ref.[17],leads to(7).Here we are interested in the additional contributions arising at orderαs.They are shown in Fig.1(a)-(c).Besides a two-loop perturbative contribution, one encounters further nonperturbative corrections proportional to the quark and the gluon condensate.Let usfirst present the result for the nonperturbative power corrections.WefindK cond(T,ω0,y)=αs ¯q q TT + αs GG y+1− ¯q g sσαβGαβq√y2−1),δn(x)=1(4π)D×1dλλ1−D∞λd u1∞1/λd u2(u1u2−1)D/2−2where C F=(N2c−1)/2N c,and D is the dimension of space-time.For D=4,the integrand diverges asλ→0.To regulate the integral,we assume D<2and use a triple integration by parts inλto obtain an expression which can be analytically continued to the vicinity of D=4.Next we set D=4+2ǫ,expand inǫ,write the result as an integral overω+,and introduce back the continuum threshold.This givesK pert(T,ω0,y)=−αsy+1 2ω0dω+ω3+e−ω+/T(16)× 12−23∂µ+3αs9π¯Λ,(17)which shows that divergences arise at orderαs.At this order,the renormalization of the sum rule is thus accomplished by a renormalization of the“bare”parameter G2in(12).In the9π¯Λ 1µ2 +O(g3s).(18)Hence a counterterm proportional to¯Λξ(y)has to be added to the bracket on the left-hand side of the sum rule(12).To evaluate its effect on the right-hand side,we note that in D dimensions[17]¯Λξ(y)F2e−2¯Λ/T=3y+1 2ω0dω+ω3+e−ω+/T(19)× 1+ǫ γE−ln4π+2lnω+−ln y+12T ξ(y) F2e−2¯Λ/T=αsy+1 2ω0dω+ω3+e−ω+/T 2lnµ6+ y r(y)−1+ln y+1According to Luke’stheorem,theuniversalfunction χ3(y )vanishes at zero recoil [10].Evaluating (20)for y =1,we thus obtain a sum rule for G 2(µ)and δΛ2.It reads G 2(µ)−¯ΛδΛ224π3ω00d ω+ω3+e −ω+/T ln µ12 +K cond (T,ω0,1),(21)where we have used that r (1)=1.Precisely this sum rule has been derived previously,starting from a two-current correlator,in Ref.[16].This provides a nontrivial check of our ing the fact that ξ(y )=[2/(y +1)]2+O (g s )according to (19),we find that the µ-dependent terms cancel out when we eliminate G 2(µ)and δΛ2from the sum rule for χ3(y ).Before we present our final result,there is one more effect which has to be taken into account,namely a spin-symmetry violating correction to the continuum threshold ω0.Since the chromo-magnetic interaction changes the masses of the ground-state mesons [cf.(10)],it also changes the masses of higher resonance states.Expanding the physical threshold asωphys =ω0 1+d M8π3 22 δ3 ω032π2ω30e −ω0/T 26π2−r (y )−ξ(y ) δ0 ω096π 248T 1−ξ(y ).It explicitly exhibits the fact that χ3(1)=0.III.NUMERICAL ANALYSISLet us now turn to the evaluation of the sum rule (23).For the QCD parameters we take the standard values¯q q =−(0.23GeV)3,αs GG =0.04GeV4,¯q g sσαβGαβq =m20 ¯q q ,m20=0.8GeV2.(24) Furthermore,we useδω2=−0.1GeV from above,andαs/π=0.1corresponding to the scale µ=2¯Λ≃1GeV,which is appropriate for evaluating radiative corrections in the effective theory[15].The sensitivity of our results to changes in these parameters will be discussed below.The dependence of the left-hand side of(23)on¯Λand F can be eliminated by using a QCD sum rule for these parameters,too.It reads[16]¯ΛF2e−2¯Λ/T=9T4T − ¯q g sσαβGαβq4π2 2T − ¯q q +(2y+1)4T2.(26) Combining(23),(25)and(26),we obtainχ3(y)as a function ofω0and T.These parameters can be determined from the analysis of a QCD sum rule for the correlator of two heavy-light currents in the effective theory[16,18].Onefinds good stability forω0=2.0±0.3GeV,and the consistency of the theoretical calculation requires that the Borel parameter be in the range0.6<T<1.0GeV.It supports the self-consistency of the approach that,as shown in Fig.2,wefind stability of the sum rule(23)in the same region of parameter space.Note that it is in fact theδω2-term that stabilizes the sum rule.Without it there were no plateau.Over the kinematic range accessible in semileptonic B→D(∗)ℓνdecays,we show in Fig.3(a)the range of predictions forχ3(y)obtained for1.7<ω0<2.3GeV and0.7<T< 1.2GeV.From this we estimate a relative uncertainty of∼±25%,which is mainly due to the uncertainty in the continuum threshold.It is apparent that the form factor is small,not exceeding the level of1%.2Finally,we show in Fig.3(b)the contributions of the individual terms in the sum rule (23).Due to the large negative contribution proportional to the quark condensate,the terms of orderαs,which we have calculated in this paper,cancel each other to a large extent.As a consequence,ourfinal result forχ3(y)is not very different from that obtained neglecting these terms[17].This is,however,an accident.For instance,the order-αs corrections would enhance the sum rule prediction by a factor of two if the ¯q q -term had the opposite sign. From thisfigure one can also deduce how changes in the values of the vacuum condensates would affect the numerical results.As long as one stays within the standard limits,the sensitivity to such changes is in fact rather small.For instance,working with the larger value ¯q q =−(0.26GeV)3,or varying m20between0.6and1.0GeV2,changesχ3(y)by no more than±0.15%.In conclusion,we have presented the complete order-αs QCD sum rule analysis of the subleading Isgur-Wise functionχ3(y),including in particular the two-loop perturbative con-tribution.Wefind that over the kinematic region accessible in semileptonic B decays this form factor is small,typically of the order of1%.When combined with our previous analysis [20],which predicted similarly small values for the universal functionχ2(y),these results strongly indicate that power corrections in the heavy quark expansion which are induced by the chromo-magnetic interaction between the gluonfield and the heavy quark spin are small.ACKNOWLEDGMENTSIt is a pleasure to thank Michael Peskin for helpful discussions.M.N.gratefully acknowl-edgesfinancial support from the BASF Aktiengesellschaft and from the German National Scholarship Foundation.Y.N.is an incumbent of the Ruth E.Recu Career Development chair,and is supported in part by the Israel Commission for Basic Research and by the Minerva Foundation.This work was also supported by the Department of Energy,contract DE-AC03-76SF00515.REFERENCES[1]E.Eichten and B.Hill,Phys.Lett.B234,511(1990);243,427(1990).[2]B.Grinstein,Nucl.Phys.B339,253(1990).[3]H.Georgi,Phys.Lett.B240,447(1990).[4]T.Mannel,W.Roberts and Z.Ryzak,Nucl.Phys.B368,204(1992).[5]A.F.Falk,H.Georgi,B.Grinstein,and M.B.Wise,Nucl.Phys.B343,1(1990).[6]N.Isgur and M.B.Wise,Phys.Lett.B232,113(1989);237,527(1990).[7]J.D.Bjorken,Proceedings of the18th SLAC Summer Institute on Particle Physics,pp.167,Stanford,California,July1990,edited by J.F.Hawthorne(SLAC,Stanford,1991).[8]M.B.Voloshin and M.A.Shifman,Yad.Fiz.45,463(1987)[Sov.J.Nucl.Phys.45,292(1987)];47,801(1988)[47,511(1988)].[9]A.F.Falk,B.Grinstein,and M.E.Luke,Nucl.Phys.B357,185(1991).[10]M.E.Luke,Phys.Lett.B252,447(1990).[11]A.F.Falk,M.Neubert,and M.E.Luke,SLAC preprint SLAC–PUB–5771(1992),toappear in Nucl.Phys.B.[12]M.Neubert,V.Rieckert,B.Stech,and Q.P.Xu,in Heavy Flavours,edited by A.J.Buras and M.Lindner,Advanced Series on Directions in High Energy Physics(World Scientific,Singapore,1992).[13]A.V.Radyushkin,Phys.Lett.B271,218(1991).[14]D.J.Broadhurst and A.G.Grozin,Phys.Lett.B274,421(1992).[15]M.Neubert,Phys.Rev.D45,2451(1992).[16]M.Neubert,Phys.Rev.D46,1076(1992).[17]M.Neubert,Phys.Rev.D46,3914(1992).[18]E.Bagan,P.Ball,V.M.Braun,and H.G.Dosch,Phys.Lett.B278,457(1992);E.Bagan,P.Ball,and P.Gosdzinsky,Heidelberg preprint HD–THEP–92–40(1992).[19]B.Blok and M.Shifman,Santa Barbara preprint NSF–ITP–92–100(1992).[20]M.Neubert,Z.Ligeti,and Y.Nir,SLAC preprint SLAC–PUB–5915(1992).[21]M.Neubert,SLAC preprint SLAC–PUB–5992(1992).[22]M.A.Shifman,A.I.Vainshtein,and V.I.Zakharov,Nucl.Phys.B147,385(1979);B147,448(1979).FIGURESFIG.1.Diagrams contributing to the sum rule for the universal form factorχ3(v·v′):two-loop perturbative contribution(a),and nonperturbative contributions proportional to the quark con-densate(b),the gluon condensate(c),and the mixed condensate(d).Heavy quark propagators are drawn as double lines.The square represents the chromo-magnetic operator.FIG.2.Analysis of the stability region for the sum rule(23):The form factorχ3(y)is shown for y=1.5as a function of the Borel parameter.From top to bottom,the solid curves refer toω0=1.7,2.0,and2.3GeV.The dashes lines are obtained by neglecting the contribution proportional toδω2.FIG.3.(a)Prediction for the form factorχ3(v·v′)in the stability region1.7<ω0<2.3 GeV and0.7<T<1.2GeV.(b)Individual contributions toχ3(v·v′)for T=0.8GeV and ω0=2.0GeV:total(solid),mixed condensate(dashed-dotted),gluon condensate(wide dots), quark condensate(dashes).The perturbative contribution and theδω2-term are indistinguishable in thisfigure and are both represented by the narrow dots.11。
DOI: 10.1126/science.1094786, 441 (2004);304Science et al.Mitchell S. Abrahamsen,Cryptosporidium parvum Complete Genome Sequence of the Apicomplexan, (this information is current as of October 7, 2009 ):The following resources related to this article are available online at/cgi/content/full/304/5669/441version of this article at:including high-resolution figures, can be found in the online Updated information and services,/cgi/content/full/1094786/DC1 can be found at:Supporting Online Material/cgi/content/full/304/5669/441#otherarticles , 9 of which can be accessed for free: cites 25 articles This article 239 article(s) on the ISI Web of Science. cited by This article has been /cgi/content/full/304/5669/441#otherarticles 53 articles hosted by HighWire Press; see: cited by This article has been/cgi/collection/genetics Genetics: subject collections This article appears in the following/about/permissions.dtl in whole or in part can be found at: this article permission to reproduce of this article or about obtaining reprints Information about obtaining registered trademark of AAAS.is a Science 2004 by the American Association for the Advancement of Science; all rights reserved. The title Copyright American Association for the Advancement of Science, 1200 New York Avenue NW, Washington, DC 20005. 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Apicomplexan,Cryptosporidium parvumMitchell S.Abrahamsen,1,2*†Thomas J.Templeton,3†Shinichiro Enomoto,1Juan E.Abrahante,1Guan Zhu,4 Cheryl ncto,1Mingqi Deng,1Chang Liu,1‡Giovanni Widmer,5Saul Tzipori,5GregoryA.Buck,6Ping Xu,6 Alan T.Bankier,7Paul H.Dear,7Bernard A.Konfortov,7 Helen F.Spriggs,7Lakshminarayan Iyer,8Vivek Anantharaman,8L.Aravind,8Vivek Kapur2,9The apicomplexan Cryptosporidium parvum is an intestinal parasite that affects healthy humans and animals,and causes an unrelenting infection in immuno-compromised individuals such as AIDS patients.We report the complete ge-nome sequence of C.parvum,type II isolate.Genome analysis identifies ex-tremely streamlined metabolic pathways and a reliance on the host for nu-trients.In contrast to Plasmodium and Toxoplasma,the parasite lacks an api-coplast and its genome,and possesses a degenerate mitochondrion that has lost its genome.Several novel classes of cell-surface and secreted proteins with a potential role in host interactions and pathogenesis were also detected.Elu-cidation of the core metabolism,including enzymes with high similarities to bacterial and plant counterparts,opens new avenues for drug development.Cryptosporidium parvum is a globally impor-tant intracellular pathogen of humans and animals.The duration of infection and patho-genesis of cryptosporidiosis depends on host immune status,ranging from a severe but self-limiting diarrhea in immunocompetent individuals to a life-threatening,prolonged infection in immunocompromised patients.Asubstantial degree of morbidity and mortalityis associated with infections in AIDS pa-tients.Despite intensive efforts over the past20years,there is currently no effective ther-apy for treating or preventing C.parvuminfection in humans.Cryptosporidium belongs to the phylumApicomplexa,whose members share a com-mon apical secretory apparatus mediating lo-comotion and tissue or cellular invasion.Many apicomplexans are of medical or vet-erinary importance,including Plasmodium,Babesia,Toxoplasma,Neosprora,Sarcocys-tis,Cyclospora,and Eimeria.The life cycle ofC.parvum is similar to that of other cyst-forming apicomplexans(e.g.,Eimeria and Tox-oplasma),resulting in the formation of oocysts1Department of Veterinary and Biomedical Science,College of Veterinary Medicine,2Biomedical Genom-ics Center,University of Minnesota,St.Paul,MN55108,USA.3Department of Microbiology and Immu-nology,Weill Medical College and Program in Immu-nology,Weill Graduate School of Medical Sciences ofCornell University,New York,NY10021,USA.4De-partment of Veterinary Pathobiology,College of Vet-erinary Medicine,Texas A&M University,College Sta-tion,TX77843,USA.5Division of Infectious Diseases,Tufts University School of Veterinary Medicine,NorthGrafton,MA01536,USA.6Center for the Study ofBiological Complexity and Department of Microbiol-ogy and Immunology,Virginia Commonwealth Uni-versity,Richmond,VA23198,USA.7MRC Laboratoryof Molecular Biology,Hills Road,Cambridge CB22QH,UK.8National Center for Biotechnology Infor-mation,National Library of Medicine,National Insti-tutes of Health,Bethesda,MD20894,USA.9Depart-ment of Microbiology,University of Minnesota,Min-neapolis,MN55455,USA.*To whom correspondence should be addressed.E-mail:abe@†These authors contributed equally to this work.‡Present address:Bioinformatics Division,Genetic Re-search,GlaxoSmithKline Pharmaceuticals,5MooreDrive,Research Triangle Park,NC27009,USA.R E P O R T S SCIENCE VOL30416APRIL2004441o n O c t o b e r 7 , 2 0 0 9 w w w . s c i e n c e m a g . o r g D o w n l o a d e d f r o mthat are shed in the feces of infected hosts.C.parvum oocysts are highly resistant to environ-mental stresses,including chlorine treatment of community water supplies;hence,the parasite is an important water-and food-borne pathogen (1).The obligate intracellular nature of the par-asite ’s life cycle and the inability to culture the parasite continuously in vitro greatly impair researchers ’ability to obtain purified samples of the different developmental stages.The par-asite cannot be genetically manipulated,and transformation methodologies are currently un-available.To begin to address these limitations,we have obtained the complete C.parvum ge-nome sequence and its predicted protein com-plement.(This whole-genome shotgun project has been deposited at DDBJ/EMBL/GenBank under the project accession AAEE00000000.The version described in this paper is the first version,AAEE01000000.)The random shotgun approach was used to obtain the complete DNA sequence (2)of the Iowa “type II ”isolate of C.parvum .This isolate readily transmits disease among numerous mammals,including humans.The resulting ge-nome sequence has roughly 13ϫgenome cov-erage containing five gaps and 9.1Mb of totalDNA sequence within eight chromosomes.The C.parvum genome is thus quite compact rela-tive to the 23-Mb,14-chromosome genome of Plasmodium falciparum (3);this size difference is predominantly the result of shorter intergenic regions,fewer introns,and a smaller number of genes (Table 1).Comparison of the assembled sequence of chromosome VI to that of the recently published sequence of chromosome VI (4)revealed that our assembly contains an ad-ditional 160kb of sequence and a single gap versus two,with the common sequences dis-playing a 99.993%sequence identity (2).The relative paucity of introns greatly simplified gene predictions and facilitated an-notation (2)of predicted open reading frames (ORFs).These analyses provided an estimate of 3807protein-encoding genes for the C.parvum genome,far fewer than the estimated 5300genes predicted for the Plasmodium genome (3).This difference is primarily due to the absence of an apicoplast and mitochondrial genome,as well as the pres-ence of fewer genes encoding metabolic functions and variant surface proteins,such as the P.falciparum var and rifin molecules (Table 2).An analysis of the encoded pro-tein sequences with the program SEG (5)shows that these protein-encoding genes are not enriched in low-complexity se-quences (34%)to the extent observed in the proteins from Plasmodium (70%).Our sequence analysis indicates that Cryptosporidium ,unlike Plasmodium and Toxoplasma ,lacks both mitochondrion and apicoplast genomes.The overall complete-ness of the genome sequence,together with the fact that similar DNA extraction proce-dures used to isolate total genomic DNA from C.parvum efficiently yielded mito-chondrion and apicoplast genomes from Ei-meria sp.and Toxoplasma (6,7),indicates that the absence of organellar genomes was unlikely to have been the result of method-ological error.These conclusions are con-sistent with the absence of nuclear genes for the DNA replication and translation machinery characteristic of mitochondria and apicoplasts,and with the lack of mito-chondrial or apicoplast targeting signals for tRNA synthetases.A number of putative mitochondrial pro-teins were identified,including components of a mitochondrial protein import apparatus,chaperones,uncoupling proteins,and solute translocators (table S1).However,the ge-nome does not encode any Krebs cycle en-zymes,nor the components constituting the mitochondrial complexes I to IV;this finding indicates that the parasite does not rely on complete oxidation and respiratory chains for synthesizing adenosine triphosphate (ATP).Similar to Plasmodium ,no orthologs for the ␥,␦,or εsubunits or the c subunit of the F 0proton channel were detected (whereas all subunits were found for a V-type ATPase).Cryptosporidium ,like Eimeria (8)and Plas-modium ,possesses a pyridine nucleotide tran-shydrogenase integral membrane protein that may couple reduced nicotinamide adenine dinucleotide (NADH)and reduced nico-tinamide adenine dinucleotide phosphate (NADPH)redox to proton translocation across the inner mitochondrial membrane.Unlike Plasmodium ,the parasite has two copies of the pyridine nucleotide transhydrogenase gene.Also present is a likely mitochondrial membrane –associated,cyanide-resistant alter-native oxidase (AOX )that catalyzes the reduction of molecular oxygen by ubiquinol to produce H 2O,but not superoxide or H 2O 2.Several genes were identified as involved in biogenesis of iron-sulfur [Fe-S]complexes with potential mitochondrial targeting signals (e.g.,nifS,nifU,frataxin,and ferredoxin),supporting the presence of a limited electron flux in the mitochondrial remnant (table S2).Our sequence analysis confirms the absence of a plastid genome (7)and,additionally,the loss of plastid-associated metabolic pathways including the type II fatty acid synthases (FASs)and isoprenoid synthetic enzymes thatTable 1.General features of the C.parvum genome and comparison with other single-celled eukaryotes.Values are derived from respective genome project summaries (3,26–28).ND,not determined.FeatureC.parvum P.falciparum S.pombe S.cerevisiae E.cuniculiSize (Mbp)9.122.912.512.5 2.5(G ϩC)content (%)3019.43638.347No.of genes 38075268492957701997Mean gene length (bp)excluding introns 1795228314261424ND Gene density (bp per gene)23824338252820881256Percent coding75.352.657.570.590Genes with introns (%)553.9435ND Intergenic regions (G ϩC)content %23.913.632.435.145Mean length (bp)5661694952515129RNAsNo.of tRNA genes 454317429944No.of 5S rRNA genes 6330100–2003No.of 5.8S ,18S ,and 28S rRNA units 57200–400100–20022Table parison between predicted C.parvum and P.falciparum proteins.FeatureC.parvum P.falciparum *Common †Total predicted proteins380752681883Mitochondrial targeted/encoded 17(0.45%)246(4.7%)15Apicoplast targeted/encoded 0581(11.0%)0var/rif/stevor ‡0236(4.5%)0Annotated as protease §50(1.3%)31(0.59%)27Annotated as transporter 69(1.8%)34(0.65%)34Assigned EC function ¶167(4.4%)389(7.4%)113Hypothetical proteins925(24.3%)3208(60.9%)126*Values indicated for P.falciparum are as reported (3)with the exception of those for proteins annotated as protease or transporter.†TBLASTN hits (e Ͻ–5)between C.parvum and P.falciparum .‡As reported in (3).§Pre-dicted proteins annotated as “protease or peptidase”for C.parvum (CryptoGenome database,)and P.falciparum (PlasmoDB database,).Predicted proteins annotated as “trans-porter,permease of P-type ATPase”for C.parvum (CryptoGenome)and P.falciparum (PlasmoDB).¶Bidirectional BLAST hit (e Ͻ–15)to orthologs with assigned Enzyme Commission (EC)numbers.Does not include EC assignment numbers for protein kinases or protein phosphatases (due to inconsistent annotation across genomes),or DNA polymerases or RNA polymerases,as a result of issues related to subunit inclusion.(For consistency,46proteins were excluded from the reported P.falciparum values.)R E P O R T S16APRIL 2004VOL 304SCIENCE 442 o n O c t o b e r 7, 2009w w w .s c i e n c e m a g .o r g D o w n l o a d e d f r o mare otherwise localized to the plastid in other apicomplexans.C.parvum fatty acid biosynthe-sis appears to be cytoplasmic,conducted by a large(8252amino acids)modular type I FAS (9)and possibly by another large enzyme that is related to the multidomain bacterial polyketide synthase(10).Comprehensive screening of the C.parvum genome sequence also did not detect orthologs of Plasmodium nuclear-encoded genes that contain apicoplast-targeting and transit sequences(11).C.parvum metabolism is greatly stream-lined relative to that of Plasmodium,and in certain ways it is reminiscent of that of another obligate eukaryotic parasite,the microsporidian Encephalitozoon.The degeneration of the mi-tochondrion and associated metabolic capabili-ties suggests that the parasite largely relies on glycolysis for energy production.The parasite is capable of uptake and catabolism of mono-sugars(e.g.,glucose and fructose)as well as synthesis,storage,and catabolism of polysac-charides such as trehalose and amylopectin. Like many anaerobic organisms,it economizes ATP through the use of pyrophosphate-dependent phosphofructokinases.The conver-sion of pyruvate to acetyl–coenzyme A(CoA) is catalyzed by an atypical pyruvate-NADPH oxidoreductase(Cp PNO)that contains an N-terminal pyruvate–ferredoxin oxidoreductase (PFO)domain fused with a C-terminal NADPH–cytochrome P450reductase domain (CPR).Such a PFO-CPR fusion has previously been observed only in the euglenozoan protist Euglena gracilis(12).Acetyl-CoA can be con-verted to malonyl-CoA,an important precursor for fatty acid and polyketide biosynthesis.Gly-colysis leads to several possible organic end products,including lactate,acetate,and ethanol. The production of acetate from acetyl-CoA may be economically beneficial to the parasite via coupling with ATP production.Ethanol is potentially produced via two in-dependent pathways:(i)from the combination of pyruvate decarboxylase and alcohol dehy-drogenase,or(ii)from acetyl-CoA by means of a bifunctional dehydrogenase(adhE)with ac-etaldehyde and alcohol dehydrogenase activi-ties;adhE first converts acetyl-CoA to acetal-dehyde and then reduces the latter to ethanol. AdhE predominantly occurs in bacteria but has recently been identified in several protozoans, including vertebrate gut parasites such as Enta-moeba and Giardia(13,14).Adjacent to the adhE gene resides a second gene encoding only the AdhE C-terminal Fe-dependent alcohol de-hydrogenase domain.This gene product may form a multisubunit complex with AdhE,or it may function as an alternative alcohol dehydro-genase that is specific to certain growth condi-tions.C.parvum has a glycerol3-phosphate dehydrogenase similar to those of plants,fungi, and the kinetoplastid Trypanosoma,but(unlike trypanosomes)the parasite lacks an ortholog of glycerol kinase and thus this pathway does not yield glycerol production.In addition to themodular fatty acid synthase(Cp FAS1)andpolyketide synthase homolog(Cp PKS1), C.parvum possesses several fatty acyl–CoA syn-thases and a fatty acyl elongase that may partici-pate in fatty acid metabolism.Further,enzymesfor the metabolism of complex lipids(e.g.,glyc-erolipid and inositol phosphate)were identified inthe genome.Fatty acids are apparently not anenergy source,because enzymes of the fatty acidoxidative pathway are absent,with the exceptionof a3-hydroxyacyl-CoA dehydrogenase.C.parvum purine metabolism is greatlysimplified,retaining only an adenosine ki-nase and enzymes catalyzing conversionsof adenosine5Ј-monophosphate(AMP)toinosine,xanthosine,and guanosine5Ј-monophosphates(IMP,XMP,and GMP).Among these enzymes,IMP dehydrogenase(IMPDH)is phylogenetically related toε-proteobacterial IMPDH and is strikinglydifferent from its counterparts in both thehost and other apicomplexans(15).In con-trast to other apicomplexans such as Toxo-plasma gondii and P.falciparum,no geneencoding hypoxanthine-xanthineguaninephosphoribosyltransferase(HXGPRT)is de-tected,in contrast to a previous report on theactivity of this enzyme in C.parvum sporo-zoites(16).The absence of HXGPRT sug-gests that the parasite may rely solely on asingle enzyme system including IMPDH toproduce GMP from AMP.In contrast to otherapicomplexans,the parasite appears to relyon adenosine for purine salvage,a modelsupported by the identification of an adeno-sine transporter.Unlike other apicomplexansand many parasitic protists that can synthe-size pyrimidines de novo,C.parvum relies onpyrimidine salvage and retains the ability forinterconversions among uridine and cytidine5Ј-monophosphates(UMP and CMP),theirdeoxy forms(dUMP and dCMP),and dAMP,as well as their corresponding di-and triphos-phonucleotides.The parasite has also largelyshed the ability to synthesize amino acids denovo,although it retains the ability to convertselect amino acids,and instead appears torely on amino acid uptake from the host bymeans of a set of at least11amino acidtransporters(table S2).Most of the Cryptosporidium core pro-cesses involved in DNA replication,repair,transcription,and translation conform to thebasic eukaryotic blueprint(2).The transcrip-tional apparatus resembles Plasmodium interms of basal transcription machinery.How-ever,a striking numerical difference is seenin the complements of two RNA bindingdomains,Sm and RRM,between P.falcipa-rum(17and71domains,respectively)and C.parvum(9and51domains).This reductionresults in part from the loss of conservedproteins belonging to the spliceosomal ma-chinery,including all genes encoding Smdomain proteins belonging to the U6spliceo-somal particle,which suggests that this par-ticle activity is degenerate or entirely lost.This reduction in spliceosomal machinery isconsistent with the reduced number of pre-dicted introns in Cryptosporidium(5%)rela-tive to Plasmodium(Ͼ50%).In addition,keycomponents of the small RNA–mediatedposttranscriptional gene silencing system aremissing,such as the RNA-dependent RNApolymerase,Argonaute,and Dicer orthologs;hence,RNA interference–related technolo-gies are unlikely to be of much value intargeted disruption of genes in C.parvum.Cryptosporidium invasion of columnarbrush border epithelial cells has been de-scribed as“intracellular,but extracytoplas-mic,”as the parasite resides on the surface ofthe intestinal epithelium but lies underneaththe host cell membrane.This niche may al-low the parasite to evade immune surveil-lance but take advantage of solute transportacross the host microvillus membrane or theextensively convoluted parasitophorous vac-uole.Indeed,Cryptosporidium has numerousgenes(table S2)encoding families of putativesugar transporters(up to9genes)and aminoacid transporters(11genes).This is in starkcontrast to Plasmodium,which has fewersugar transporters and only one putative ami-no acid transporter(GenBank identificationnumber23612372).As a first step toward identification ofmulti–drug-resistant pumps,the genome se-quence was analyzed for all occurrences ofgenes encoding multitransmembrane proteins.Notable are a set of four paralogous proteinsthat belong to the sbmA family(table S2)thatare involved in the transport of peptide antibi-otics in bacteria.A putative ortholog of thePlasmodium chloroquine resistance–linkedgene Pf CRT(17)was also identified,althoughthe parasite does not possess a food vacuole likethe one seen in Plasmodium.Unlike Plasmodium,C.parvum does notpossess extensive subtelomeric clusters of anti-genically variant proteins(exemplified by thelarge families of var and rif/stevor genes)thatare involved in immune evasion.In contrast,more than20genes were identified that encodemucin-like proteins(18,19)having hallmarksof extensive Thr or Ser stretches suggestive ofglycosylation and signal peptide sequences sug-gesting secretion(table S2).One notable exam-ple is an11,700–amino acid protein with anuninterrupted stretch of308Thr residues(cgd3_720).Although large families of secretedproteins analogous to the Plasmodium multi-gene families were not found,several smallermultigene clusters were observed that encodepredicted secreted proteins,with no detectablesimilarity to proteins from other organisms(Fig.1,A and B).Within this group,at leastfour distinct families appear to have emergedthrough gene expansions specific to the Cryp-R E P O R T S SCIENCE VOL30416APRIL2004443o n O c t o b e r 7 , 2 0 0 9 w w w . s c i e n c e m a g . o r g D o w n l o a d e d f r o mtosporidium clade.These families —SKSR,MEDLE,WYLE,FGLN,and GGC —were named after well-conserved sequence motifs (table S2).Reverse transcription polymerase chain reaction (RT-PCR)expression analysis (20)of one cluster,a locus of seven adjacent CpLSP genes (Fig.1B),shows coexpression during the course of in vitro development (Fig.1C).An additional eight genes were identified that encode proteins having a periodic cysteine structure similar to the Cryptosporidium oocyst wall protein;these eight genes are similarly expressed during the onset of oocyst formation and likely participate in the formation of the coccidian rigid oocyst wall in both Cryptospo-ridium and Toxoplasma (21).Whereas the extracellular proteins described above are of apparent apicomplexan or lineage-specific in-vention,Cryptosporidium possesses many genesencodingsecretedproteinshavinglineage-specific multidomain architectures composed of animal-and bacterial-like extracellular adhe-sive domains (fig.S1).Lineage-specific expansions were ob-served for several proteases (table S2),in-cluding an aspartyl protease (six genes),a subtilisin-like protease,a cryptopain-like cys-teine protease (five genes),and a Plas-modium falcilysin-like (insulin degrading enzyme –like)protease (19genes).Nine of the Cryptosporidium falcilysin genes lack the Zn-chelating “HXXEH ”active site motif and are likely to be catalytically inactive copies that may have been reused for specific protein-protein interactions on the cell sur-face.In contrast to the Plasmodium falcilysin,the Cryptosporidium genes possess signal peptide sequences and are likely trafficked to a secretory pathway.The expansion of this family suggests either that the proteins have distinct cleavage specificities or that their diversity may be related to evasion of a host immune response.Completion of the C.parvum genome se-quence has highlighted the lack of conven-tional drug targets currently pursued for the control and treatment of other parasitic protists.On the basis of molecular and bio-chemical studies and drug screening of other apicomplexans,several putative Cryptospo-ridium metabolic pathways or enzymes have been erroneously proposed to be potential drug targets (22),including the apicoplast and its associated metabolic pathways,the shikimate pathway,the mannitol cycle,the electron transport chain,and HXGPRT.Nonetheless,complete genome sequence analysis identifies a number of classic and novel molecular candidates for drug explora-tion,including numerous plant-like and bacterial-like enzymes (tables S3and S4).Although the C.parvum genome lacks HXGPRT,a potent drug target in other api-complexans,it has only the single pathway dependent on IMPDH to convert AMP to GMP.The bacterial-type IMPDH may be a promising target because it differs substan-tially from that of eukaryotic enzymes (15).Because of the lack of de novo biosynthetic capacity for purines,pyrimidines,and amino acids,C.parvum relies solely on scavenge from the host via a series of transporters,which may be exploited for chemotherapy.C.parvum possesses a bacterial-type thymidine kinase,and the role of this enzyme in pyrim-idine metabolism and its drug target candida-cy should be pursued.The presence of an alternative oxidase,likely targeted to the remnant mitochondrion,gives promise to the study of salicylhydroxamic acid (SHAM),as-cofuranone,and their analogs as inhibitors of energy metabolism in the parasite (23).Cryptosporidium possesses at least 15“plant-like ”enzymes that are either absent in or highly divergent from those typically found in mammals (table S3).Within the glycolytic pathway,the plant-like PPi-PFK has been shown to be a potential target in other parasites including T.gondii ,and PEPCL and PGI ap-pear to be plant-type enzymes in C.parvum .Another example is a trehalose-6-phosphate synthase/phosphatase catalyzing trehalose bio-synthesis from glucose-6-phosphate and uridine diphosphate –glucose.Trehalose may serve as a sugar storage source or may function as an antidesiccant,antioxidant,or protein stability agent in oocysts,playing a role similar to that of mannitol in Eimeria oocysts (24).Orthologs of putative Eimeria mannitol synthesis enzymes were not found.However,two oxidoreductases (table S2)were identified in C.parvum ,one of which belongs to the same families as the plant mannose dehydrogenases (25)and the other to the plant cinnamyl alcohol dehydrogenases.In principle,these enzymes could synthesize protective polyol compounds,and the former enzyme could use host-derived mannose to syn-thesize mannitol.References and Notes1.D.G.Korich et al .,Appl.Environ.Microbiol.56,1423(1990).2.See supportingdata on Science Online.3.M.J.Gardner et al .,Nature 419,498(2002).4.A.T.Bankier et al .,Genome Res.13,1787(2003).5.J.C.Wootton,Comput.Chem.18,269(1994).Fig.1.(A )Schematic showing the chromosomal locations of clusters of potentially secreted proteins.Numbers of adjacent genes are indicated in paren-theses.Arrows indicate direc-tion of clusters containinguni-directional genes (encoded on the same strand);squares indi-cate clusters containingg enes encoded on both strands.Non-paralogous genes are indicated by solid gray squares or direc-tional triangles;SKSR (green triangles),FGLN (red trian-gles),and MEDLE (blue trian-gles)indicate three C.parvum –specific families of paralogous genes predominantly located at telomeres.Insl (yellow tri-angles)indicates an insulinase/falcilysin-like paralogous gene family.Cp LSP (white square)indicates the location of a clus-ter of adjacent large secreted proteins (table S2)that are cotranscriptionally regulated.Identified anchored telomeric repeat sequences are indicated by circles.(B )Schematic show-inga select locus containinga cluster of coexpressed large secreted proteins (Cp LSP).Genes and intergenic regions (regions between identified genes)are drawn to scale at the nucleotide level.The length of the intergenic re-gions is indicated above or be-low the locus.(C )Relative ex-pression levels of CpLSP (red lines)and,as a control,C.parvum Hedgehog-type HINT domain gene (blue line)duringin vitro development,as determined by semiquantitative RT-PCR usingg ene-specific primers correspondingto the seven adjacent g enes within the CpLSP locus as shown in (B).Expression levels from three independent time-course experiments are represented as the ratio of the expression of each gene to that of C.parvum 18S rRNA present in each of the infected samples (20).R E P O R T S16APRIL 2004VOL 304SCIENCE 444 o n O c t o b e r 7, 2009w w w .s c i e n c e m a g .o r g D o w n l o a d e d f r o m。
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Journal of Finance, 71(1).[3]Chow, V., & Lai, C. W. (2015). Conditional sharpe ratios. Finance Research Letters, 12, 117-133.海外文献推荐-第十一期参考文献：[1] Mladina, P. (2017). Illuminating hedge fund returns to improve portfolio construction. Social Science Electronic Publishing, 41(3), 127-139.[2] Choi, N., Fedenia, M., Skiba, H., & Sokolyk, T. (2016). Portfolio concentration and performance of institutional investors worldwide. Journal of Financial Economics.[3] Martijn Boons, 2016, State variables, macroeconomic activity, and the cross section of individual stocks, Journal of Financial Economics 119, 489-511海外文献推荐-第十二期参考文献：[1] Blanchett, D., & Ratner, H. (2015). Building efficient income portfolios. Journal of Portfolio Management, 41(3), 117-125.[2] Özde Öztekin. (2015). Capital structure decisions around the world: which factors are reliably important?. Journal of Financial & Quantitative Analysis, 50(3).[3] 2015, Does the number of stocks in a portfolio influence performance? 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ResearchGate海外文献推荐-第七十八期参考文献：[1] Joel Hasbrouck. High-Frequency Quoting: Short-Term V olatility in Bids and Offers. JOURNAL OF FINANCIAL AND QUANTITATIVE ANALYSIS海外文献推荐-第七十九期参考文献：[1] Tarun Gupta and Bryan Kelly. Factor Momentum Everywhere. Institutional Investor Journals海外文献推荐-第八十期参考文献：[1] MICHAEL A. BABYAK , P H D. What You See May Not Be What You Get: A Brief, Nontechnical Introduction to Overfitting in Regression-Type Models. S T A T I S T I C A L C O R N E R海外文献推荐-第八十一期参考文献：[1] Eric Jondeau , Qunzi Zhang , Xiaoneng Zhu. Average Skewness Matters.海外文献推荐-第八十二期参考文献：[1] JOHN A. HASLEM. Morningstar Mutual Fund Measures and Selection Model. THE JOURNAL OF WEALTH MANAGEMENT海外文献推荐-第八十三期参考文献：[1] EUGENE F. FAMA and KENNETH R. FRENCH. Luck versus Skill in the Cross-Section of Mutual Fund Returns. THE JOURNAL OF FINANCE海外文献推荐-第八十四期参考文献：[1] How Transparent Are ETFs?[2] Lara Crigger. Nontransparent Active: Next ETF Revolution?.海外文献推荐-第八十五期参考文献：[1] Olivier Rousse and Benoît Sévi. 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