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Suppression of cact lethality by mutations in Tl, tub and pll predicts that mutations in these latter genes should also cause hematopoietic defects. To determine if there are any obvious defects in the morphology of mutant lymph glands, we examined glands from Tl, tub and pll larvae and did not find any apparent or striking differences in the size or the organization of lobes or pericardial cells results not shown ; . We, therefore, performed hemocyte counts on Tl, tub, pll and tub pll mutants. Heterozygous siblings and wild-type larvae served as controls. As can be seen from Fig. 3, the concentration of hemocytes in these mutant combinations ranged from 21% to 48% of the wild-type concentration. Hemocyte concentration in Tl- larvae is the lowest 1, 222 cells l, as compared to 3, 771 cells l of the heterozygous control hemolymph ; . In contrast, the hemocyte concentration in the TlD background is roughly two-fold 10, 348 cells l ; compared to the Canton-S hemolymph 5, 523 cells l ; . When compared with the heterozygous controls, hemocyte concentration per microliter of tub 1, 375 versus 6, 381 ; , pll 2, 508 versus 5, 142 ; or tub pll 1, 875 versus 5, 182 ; hemolymph also shows a statistically significant deficit Fig. 3 ; . In control experiments, we found that hemocyte concentration in unrelated semilethal lethal strains is in the wild-type range R. Sorrentino and S. G., unpublished data ; , suggesting that the effects of Tl, pll and tub mutations on hemocyte counts are particular to these mutations. Since Tl, tub, and pll have phenotypic effects that are opposite of cact, a deficit in hemocyte concentration in these three `dorsal group' mutants is consistent with the genetic interaction data presented above. Together, these observations support the idea that a signal transduction pathway with Toll, Tube, Pelle and Cactus proteins is involved in controlling hemocyte density in the larval hemolymph. Mutations in Toll and cactus affect the number of cells undergoing division To investigate the cause of aberrant hemocyte concentrations in Tl and cact larvae, we examined if the number of hemocytes
Consistent with our predicted role of MCL in innate immune responses in the clam. Broad specificity of innate immune-related lectins may be a general biological phenomenon. A previous hapten inhibition assay showed that lipopolysaccharides derived from bacteria-infecting marine organisms but not lipopolysaccharides from other sources are potent inhibitors of MCL 26 ; . The tachylectin-2 from horseshoe crab binds to both D-GlcNAc and D-GalNAc and lipopolysaccharide from Gram-negative bacteria 42 ; . The C-type lectin perlucin from the abalone has been reported to have broad binding specificity with the ability to bind glycoproteins containing galactose or mannose glucose 46 ; . In the absence of immunoglobulin molecules, broad binding specificity of lectin may be required for effective humoral non-self-recognition in FIGURE 11. Immunohistochemical staining of MCL in Perkinsus-infected clam tissue. Perkinsus-infected the marine invertebrate. clams were embedded in freezing medium. Cryostat sections were incubated with pre-immune rabbit IgG A MCL can act as an opsonin to and B ; or anti-MCL rabbit antibody C and D ; and stained FITC-labeled anti-rabbit goat antibody as described under "Materials and Methods." , Perkinsus trophozoite; arrows, encapsulated bodies. stimulate in vitro phagocytosis Fig. 10, A and B ; . We also made the cessed MCL data not shown ; . It is still uncertain where the novel observation that MCL synthesis is induced by Perkinsus conversion of the 74-kDa species to the 30- and 34-kDa species infection of clams. It has been reported that many lectins from is occurring, either in the hemolymph or in a terminal destina- marine invertebrates show antibacterial activity. The lectin tion such as tissue or surface of pathogens. Future studies will from the horse mussel M. modiolus 21 ; exhibited strong antiaddress this issue. bacterial activity against tested Vibrio strains. The gigalin H and We reported earlier that mucin types 1 and 2, which contain gigalin E lectins from oyster Crassostrea gigas ; acted as opsoGalNAc residues, exhibited the best inhibition of MCL-medi- nins to stimulate in vitro phagocytosis of the marine bacterium ated hemagglutination and that highly branched mannan struc- Vibrio anguillarum 47 ; . However, the induction of lectin tures from marine-originated bacteria also showed strong inhi- involvement in defense against parasitic invasion, as we have bition 26 ; . However, because these glycan structures are not seen for Perkinsus, has not been reported previously. Our preswell characterized, and high branched mannan from Escheent study shows that only the hemocyte from Perkinsus-inrichia coli did not inhibit hemagglutination of MCL, the results fected clams produces MCL and that MCL is able to bind to the were not conclusive whether the general high mannose strucPerkinsus Figs. 2 and 8 ; . However, Perkinsus trophozoite is ture inhibited agglutination. In glycan array analyses MCL showed the highest binding to glycans containing the Lacdi- probably too large to allow direct engulfment by hemocytes. NAc motif GalNAc 1 4GlcNAc 1-R ; and to glycans with MCL may function to recruit hemocytes around Perkinsus terminal -Gal and -GalNAc structures. Many lectins from and or function in phagocytosis of parasite-derived material. marine invertebrates have affinity toward to GalNAc residues The infiltration of hemocytes Fig. 11 ; around Perkinsus is a 45 ; Interestingly, a mannose-binding lectin-like lectin from well known symptom of Perkinsus-infected clam in the gill 50 ; . invertebrates has also been reported to have binding specificity Clusters of Perkinsus trophozoites were observed encapsulated toward to GalNAc-containing glycans instead of mannose 48 ; . in amorphous eosinophilic material, forming cyst-like strucThus, the recognition of LacdiNAc-type glycans, which are tures Fig. 11 ; 50 ; . Parasite encapsulation is well characterized abundantly expressed in many invertebrates and parasites, is in insects 51 ; . For example, the C-type lectin immulectin has.
Total hemocyte counting
Turnbull MW, Martin SB, Webb BA. 2004. Quantitative analysis of hemocyte morphological abnormalities associated with Campoletis sonorensis parasitization. 15pp. Journal of Insect Science, 4: 11, Available online: insectscience 4.11.
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The objective of this study was to compare the hemocyte characteristics - morphological and functional - of oysters from six bi-parental families reared at three different experimental sites along the French coast during summer and to answer two main questions. The first question addressed was, "do oyster hemocyte characteristics have a genetic basis, independent of environmental conditions and gametogenic cycle?" To address this question, hemocyte characteristics of the six bi-parental families in this experiment were compared, regardless of the rearing site. This analysis showed that 7 of the 15 hemocyte variables differentiated at least one family from others of the 6 analyzed. Moreover, only 3 of 15 variables showed a significant interaction between site and family. ROS production was the most notably different between families under all incubation conditions. Thus, a genetic bases determining hemocyte characteristics of oysters, and especially for ROS production, is implied. The variations in ROS production observed may result from variations in activities of the enzymes involved in the production NADPH-oxidase, NO synthase ; or in the detoxification superoxide dismutase, catalase, gluthation peroxidase ; of these molecules. Indeed, the genetic basis for the ROS-detoxification mechanism has been recently reported in the mollusk Biomphalaria glabrata Goodall et al., 2004; Zelck, Von Janowsky, 2004 ; and currently is being investigated with oysters in our laboratory. Expression of genes involved in ROS detoxification has been shown to be modulated in oysters by environmental factors, such as hypoxia and PAH exposures Boutet et al., 2004; David et al., 2005 ; . The second question addressed was, "does selection of families for high or low survival to summer mortality lead to different hemocyte characteristics" and, if so "could these different hemocyte characteristics explain the observed differences of survival?" Although the number of resistant high-survival ; and susceptible low- survival ; families was low three in each group ; , six of the 15 hemocyte variables differentiated the two groups statistically. Moreover, ANOVA results showed few significant interactions between site and survival group resistant susceptible ; . This means that, except for the adhesion test, resistant and susceptible family groups respond in the same way regardless of the site. Thus, genetic selection based on survival performance can change hemocyte characteristics of C. gigas. A more fundamental question was whether or not high survival during the summer mortality period could be attributed to differences in hemocytes: the hemocyte profile of resistant oysters was compared to that of susceptible oyster families. High-survival families had higher hemocyte counts, lower inhibition of the ROS production by the Vibrio sp. S322, and lower susceptibility to bacteria in the adhesion test. These hemocyte attributes could be hypothesized to confer better capacity for high survival when challenged by pathogenic bacteria, for example. Resistant oysters, however, also had smaller granulocytes and lower ROS production. The latter parameter has been observed to be modulated by acute stress Lacoste et al., 2002 ; , infection by V. aestuarianus Labreuche et al., 2006b; Labreuche et al., 2006a ; , and dietary changes Delaporte et al., in press ; . Cellular ROS content also depends on ROS detoxification capacities and respiration processes in other organisms Sheehan, Power, 1999; Batandier et al., 2002; Cadenas, 2004; Keller et al., 2004; Manduzio et al., 2005 ; . When suppression-substractive hydridization SSH ; techniques were used during the MOREST program Huvet et al., 2004 ; , resistant and susceptible oysters were shown to express differentially a cavortin-like gene, later characterized as an hemocyte Cu Zn superoxide dismutase SOD ; gene Gonzalez et al., 2005 ; . In their experiments, these authors showed that resistant oysters had a higher Cu Zn SOD gene expression than susceptible oysters. Accordingly, gene expression is thought to contribute to ROS detoxification; this may partially explained the difference in ROS production in our experiment between resistant and susceptible oysters. Furthermore, one may speculate that selection for one or more enzymes involved in ROS production and or detoxification may have concurrently occurred with selection for summer mortality survival. Nevertheless, we call attention to the fact that hemocyte measurements were analyzed for the survivors of the selected families after the summer mortality event; the mean mortality of susceptible oysters was approximately 30 %, and 5% in resistant families. In addition, the criterion for selection was survival during the first summer, without knowledge of the exact cause of mortality. Thus, it is important to remain prudent when trying to link hemocyte characteristics with oyster survival during the period of summer mortality. Genetic selection, however, does not overcome all environmental influences. Indeed, this study demonstrates that rearing site strongly affected the hemocyte characteristics of all families of oysters and hepsera.
Hemocyte plus iron
Spinigerin against most of the bacteria tested Table II ; . Spinigerin, when assayed against filamentous fungi, showed equivalent activity to that of Aedes cecropin and PGLa. Against the yeast strain C. albicans, spinigerin was more efficient than the two -helical peptides used as references. The data presented here establish that a member of the insect order of Isoptera produces a set of antimicrobial peptides in response to a septic injury. Antimicrobial peptides are key components of innate immunity against bacteria and fungi in invertebrates, vertebrates and plants. In insects, septic injury leads to rapid synthesis of these peptides by the fat body and certain blood cells, followed by their release into the hemolymph. However, in our study on the isopteran P. spiniger, the antimicrobial molecules appear to be constitutively present. A control experiment processed with unchallenged animals gave no indication that injection of bacteria modified the spectrum and or the level of the antimicrobial compounds. Furthermore, no antimicrobial activity had been detected in the hemolymph collected from naive and immune challenged aminals. In fact, the antifungal peptide termicin was detected in hemocyte granules and in salivary glands prepared from.
Asiatica is the predominance of one cell type, namely the granular hemocyte. A characteristic feature of this hemocyte is the nature of granular inclusions, which and herceptin!
SYNOPSIS. Because tunicates rely on innate immunity, their hemocytes are important contributors to host defense. Styela clava, a solitary ascidian, have eight hemocyte subtypes. Extracts of their total hemocyte population contained multiple small 24 kDa ; antimicrobial peptides. When purified, these fell into two distinct families that were named styelins and clavanins. Styelins A-E are phenylalanine-rich, 32 residue peptides with activity against marine bacteria and human pathogens. They show considerable sequence homology to pleurocidins, antimicrobial peptides of the flounder, Pseudopleuronectes americanus. Styelin D, one of the five styelins identified by peptide isolation and cDNA cloning, was remarkable in containing 12 post-translationally modified residues, including a 6-bromotryptophan, two monohydroxylysines, four 3, 4-dihydroxyphenylalanines DOPA ; , four dihydroxylysines and one dihydroxyarginine. These modifications enhanced Styelin D's bactericidal ability at acidic pH and high salinity. A novel histochemical stain for DOPA suggested that Styelin D was restricted to granulocytes. Clavanins A-E are histidine-rich, 23 residue peptides that are C-terminally amidated and most effective at acidic pH. Clavaspirin is a newly described family member that also has potent cytotoxic properties. By immunocytochemistry, clavanins were identified in the granules of five eosinophilic granulocyte subtypes and in macrophage cytoplasm. Transmission and scanning electron micrographs of methicillin-resistant Staphylococcus aureus MRSA ; and E. coli that had been treated with Styelin D and clavaspirin suggested that both peptides induced osmotic disregulation. Treated bacteria manifested cytoplasmic swelling and extrusion of cytoplasmic contents through their peptidoglycan cell wall. The diverse array of antimicrobial peptides in S. clava hemocytes constitutes an effective host defense mechanism.
Hemocyte differentiation
We thank Dr. Benjamin Neel Beth Israel Deaconess Medical Center ; for the use of the Odyssey Infrared Imaging System. GRANTS This work was supported by National Institute of Dental and Craniofacial Research Grant DE-10877. REFERENCES 1. Aizman O, Uhlen P, Lal M, Brismar H, and Aperia A. Ouabain, a steroid hormone that signals with slow calcium oscillations. Proc Natl Acad Sci USA 98: 13420 13424, Al-Khalili L, Kotova O, Tsuchida H, Ehren I, Feraille E, Krook A, and Chibalin AV. ERK1 2 mediates insulin stimulation of Na, K-ATPase by phosphorylation of the -subunit in human skeletal muscle cells. J Biol Chem 279: 2521125218, 2004. Benes C and Soltoff SP. Modulation of PKC tyrosine phosphorylation and activity in salivary and PC-12 cells by Src kinases. J Physiol Cell Physiol 280: C1498 C1510, 2001. 4. Blanco G and Mercer RW. Isozymes of the Na-K-ATPase: heterogeneity in structure, diversity in function. J Physiol Renal Physiol 275: F633F650, 1998. 5. Cook DI, van Lennep EW, Roberts ML, and Young JA. Secretion by the major salivary glands. In: Physiology of the Gastrointestinal Tract 3rd ed. ; , edited by Johnson RL. New York: Raven, 1994, p. 10611117. 6. Dmitrieva RI and Doris PA. Ouabain is a potent promoter of growth and activator of ERK1 2 in ouabain-resistant rat renal epithelial cells. J Biol Chem 278: 28160 28166, Falin R, Veizis IE, and Cotton CU. A role for ERK1 2 in EGF- and ATP-dependent regulation of amiloride-sensitive sodium absorption. J Physiol Cell Physiol 288: C1003C1011, 2005. 8. Fiske CH and Subbarow Y. The colorimetric determination of phosphorus. J Biol Chem 66: 375 400, Guerrero C, Lecuona E, Pesce L, Ridge KM, and Sznajder JI. Dopamine regulates Na-K-ATPase in alveolar epithelial cells via MAPKERK dependent mechanisms. J Physiol Lung Cell Mol Physiol 281: L79 L85, 2001. 10. Haas M, Askari A, and Xie Z. Involvement of Src and epidermal growth factor receptor in the signal transducing function of Na K -ATPase. J Biol Chem 275: 2783227837, 2000. Haas M, Wang H, Tian J, and Xie Z. Src-mediated inter-receptor cross-talk between the Na K -ATPase and the epidermal growth factor receptor relays the signal from ouabain to mitogen-activated protein kinase. J Biol Chem 277: 18694 18702, Hoffert JD, Leitch V, Agre P, and King LS. Hypertonic induction of aquaporin-5 expression through an ERK-dependent pathway. J Biol Chem 275: 9070 9077, Khundmiri SJ and Lederer E. PTH and DA regulate Na, K-ATPase through divergent mechanisms. J Physiol Renal Physiol 282: F512 F522, 2002. 14. Kreydiyyeh SI and Al-Sadi R. The signal transduction pathway that mediates the effect of interleukin-1 on the Na -K -ATPase in LLC-PK1 cells. Pflugers Arch 448: 231238, 2004. Liu J, Tian J, Haas M, Shapiro JI, Askari A, and Xie Z. Ouabain interaction with cardiac Na K -ATPase initiates signal cascades independent of changes in intracellular Na and Ca2 concentrations. J Biol Chem 275: 27838 27844, Liu L, Abramowitz J, Askari A, and Allen JC. Role of caveolae in ouabain-induced proliferation of cultured vascular smooth muscle cells of the synthetic phenotype. J Physiol Heart Circ Physiol 287: H2173 H2182, 2004. 17. Liu L, Mohammadi K, Aynafshar B, Wang H, Li D, Liu J, Ivanov AV, Xie Z, and Askari A. Role of caveolae in signal-transducing function of cardiac Na K -ATPase. J Physiol Cell Physiol 284: C1550 C1560, 2003 and hms.
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The four major nutritional problems identifiedin Myanmarinclude: PEM WA below -2SD ; with a prevalence 28% among children under three of years, IDD with a goitreprevalence rate of 25% amongchildrenaged 6-11 years IDA prevalence 58% Hb level ; lessthan 11g dl ; among pregnant of women and VAD Bitot's spotsamongunderfives0.38% ; ranging from 0-6% in differentpartsof the country. Interventions against PEMincludethe NationalCrowth and Promotion Programme, which hasan intensified nutritioneducation component and is supported urbannutritionprogrammes well as hospital-based by as nutrition centres. Maternalnutritionmonitoring periodically is conducted antenatal at clinicsand other vital relatednutritioncomponents integrated are into the programmes the prevention for and controlof PEM.High-potency vitaminA supplementation all underfivechildren is given. As part of long-term to strategies, nation-widenutrition educationactivities and home gardening programmes 10 townships Myanmarare in progress. in in The Ministryof Education is intensifying education efforts for the prevention of noncommunicable diseases especially reducing consumption daily by fat in diets. Thrustis beingprovided PEMresearch to throughstudies household on HH ; and individual food security which is beingtakenup as a priorityissue. Studying caringbehaviour households underwo , the outcomes which in is of are to be used for programme and policy improvement HH nutrition for security.Quantitative and qualitative approaches the community have in beenusedto obtainthe requiredinformation. goitreprevalence A survey has.
Committee on Practice and Ambulatory Medicine et al. 2003 IV; Prasad and Chudley 2002; Green 1994 IV; AAP 2001; AAP 1998 CPG; American Academy of Audiology et al. 2000 CPG Applicable Indicator Strength of Indicator Type Function Modality Evidence Age, y Number D. Developmental and Behavioural Assessment 4748 For children between ages 0 and 3, a formal developmental evaluation should be performed at least: 02 47 Once between 9 and 15 mo corrected age Pr D T Once between 21 and 30 mo corrected age Pr D T For children between ages 0 and 3, a formal developmental A D R evaluation should be performed within 2 mo of suspect or abnormal developmental screening test e.g., abnormal Bayley Infant Neurodevelopment Screener ; 5055 In the absence of a formal developmental evaluation, the presence or absence of parental concerns and a multidimensional developmental screening test should be documented using standardized instruments at least: 01 50 Once during the first 6 mo Pr Once during the second 6 mo Pr Once between ages 1 and 2 Pr S Once between ages 2 and 3 Pr S Once between ages 3 and 4 Pr S Once between ages 4 and 5 Pr S 5660 A structured, age-appropriate neuromotor assessment should be performed by corrected age at least: 01 56 Once during the first 6 mo Pr Once during the second 6 mo Pr Once between ages 1 and 2 Pr S Once between ages 2 and 3 Pr S Once between ages 4 and 5 Pr S the structured neuromotor examination or the formal A F R developmental evaluation is suspect or abnormal, then a specific intervention watchful waiting with re-evaluation, specialist consultation, or a specific intervention program ; should be started within 2 mo of the suspect or abnormal finding. 06 62 If professional who is performing the neuromotor examination C Tr I recommends physical therapy or occupational therapy for the patient, then interventions should be started within 2 mo of the recommendation. F R 2 parents express concerns about their child's behaviour, then a C specific intervention watchful waiting with re-evaluation, primary care management, referral to a specialist, or referral to a specific intervention program ; should be started within 2 mo and humalog.
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Total hemocyte concentrations in hemolymph of juvenile oysters in our study ranged from 1.3 x 105 to 1.2 x IO4 cells.ml-', somewhat lower on average than in reports for adult oysters and clams, where circulating blood cell densities were typically in the
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That the lobsters were not all at the same quirement for 3atty acids. The poor f molt stage resulted in large variances results obtained with the HCO would re within dietary treatment. Space limitations sult from the lack of either the w6 or w3 prevented the use of large enough num series. That the positive effects of CLO bers of adult lobsters to overcome the molt were due to the EFA rather than some cycle variances. However, by considering other lipid component is supported by the several of the analysis as well as the molt fact that the sterol free SF-CLO ; diet incidence together, certain conclusions re also gave better growth than either CO or lating to the nutritional value of dietary HCO diets. lipids were possible. In a three month feeding study with Lobsters have a relatively low require juvenile lobsters Castell et al., 8 ; showed ment for dietary lipid, with 5% CLO in the a requirement for 0.5% of the dry weight diet giving near optimal results. The 5% of the diet as cholesterol for growth and CLO diet resulted in a significant increase survival. Though feeding adult lobsters a sterol-free diet for 10 months did result in in number of molts, weight gain and a con siderable improvement in all condition in a decrease in serum protein and calcium dices. Though higher levels of CLO did not levels, these effects of the sterol deficiency improve the mean percent weight gain, were not nearly as dramatic as those with molt, feed conversion or hemocyte counts, juvenile lobsters. The addition of \% cho lesterol to the sterol free diet SA-CLO ; there did appear to be some nitrogen spar ing effect as seen in the slight though not resulted in increased serum protein levels significant elevation of the protein content but reduced serum calcium level even more of the serum and percent edible meat of than the SF-CLO diet. It is possible the lobsters fed the diet with the higher that some other non-saponifiable compo levels of CLO. This nitrogen sparing effect nent of the CLO besides cholesterol plays of extra dietary energy supplied by the a significant role in the calcium metabolism higher lipid levels might reduce the high in the adult lobster. One possible candidate requirement for protein previously re is vitamin D. ported for adult lobsters 6 ; . The optimal LITERATURE CITED growth effect of 5% CLO is probably re lated to the essential fatty acid EFA ; 1. Herrick, F. H. 1896 ; The American Lob ster: A study of its habits and development. requirement of lobsters. Bull. U.S. Fish. Comm. 15, 1-252. The differences noted between CLO, 2. Hughes, F. H. & Matthiessen, G. C. 1962 ; CO and HCO are also probably related to Observations on the biology of the American lobster, Homarus americanas. Limnol. the EFA requirements of lobsters. Corn oil, Oceanogr. 7 3 ; , 414-421. which is high in linoleic acid wo ; 5 and 3. Stewart, J. E., Cornick, J. W. & Dingle, J. R. low in linolenic acid w3 ; , depressed 1967A ; An electronic method for counting growth in several marine or aquatic ani lobster Homarus americanus, Milne Ed mals such as rainbow trout, Salmo gairdneri wards ; hemocytes and the influence of diet on hemocyte numbers and hemolymph pro 21 ; and shrimp, Penaeus duorarum 10 ; . teins. Can. J. Zool. 45, 291-304. Cod liver oil CLO ; is high in fatty acids 4. McLeese, D. W. 1972A ; Initial experi of the o3series 13 ; . These types of fatty ments on growth of the American lobster in acids have been shown to be essential in captivity. Fish. Res. Bd. Canada, Technical Report No. 320, 1-22. the diet of rainbow trout 22 ; . Both shrimp 5. McLeese, D. W. 1972B ; Effect of several 10 ; and catfish 23 ; exhibited growth en factors on the growth of the American lobster hancement with dietary lipids high in 3 Homarus americanus ; in captivity. J. Fish. Res. Bd. Canada 29, 1725-1730. fatty acids. Shewbart and Mies 11 ; ob 6. Stewart, J. E., Cornick, J. W., Foley, D. M., tained growth enhancement in brown Li, M. F. & Bishop, C. M. 1967B ; Muscle shrimp, Penaeus aztecus, when their puri weight relationship to serum proteins, hemo fied diet was supplemented with pure cytes and hepatopancreas in the lobster linolenic acid. Thus, some marine crus Homarus americanus ; . J. Fish. Res. Bd. Canada 24, 2339-2354. taceans including two species of shrimp 7. Castell, J. D. & Budson, S. D. 1974 ; and lobsters ; appear to have an EFA reThew6 fatty acids are essential most mammals 24 ; . for rats, pigs and and hemocyte.
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