099_1_117_herz.pdf

(XU - (QWRPRO  ± 
&RPSDULVRQ RI WKH IDW DOORFDWLRQ SDWWHUQV LQ IHPDOH SLQH VDZIOLHV
+\PHQRSWHUD 'LSULRQLGDH
/HKUVWXKO IU $QJHZDQGWH =RRORJLH )RUVWZLVVHQVFKDIWOLFKH )DNXOWlW 7HFKQLVFKH 8QLYHUVLWlW 0QFKHQ $P +RFKDQJHU  ' )UHLVLQJ *HUPDQ\ HPDLO DKHU]#IUHHQHWGH .H\ ZRUGV 'LSULRQLGDH SLQH GHIROLDWRU UHVRXUFH DOORFDWLRQ RXWEUHDN WHQGHQF\ IHHGLQJ EHKDYLRXU RYLSRVLWLRQ EHKDYLRXU
$EVWUDFW 7KH DOORFDWLRQ RI IDW UHVRXUFHV WR VRPDWLF DQG UHSURGXFWLYH WLVVXHV ZDV VWXGLHG LQ IHPDOHV RI ILYH VSHFLHV RI SLQH VDZIOLHV
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ƒ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
PRELOLW\ DQG ORQJHYLW\ RI HJJOD\LQJ IHPDOHV DQG EH DVVRFLDWHG 'LSULRQLG IHPDOHV DUH SURRYLJHQLF DQG FRQWDLQ WKHLU HQWLUH ZLWK D GLVVLPLODU LQYHVWPHQW RI QXWULHQW UHVRXUFHV LQ UHSURGXF VXSSO\ RI PDWXUH HJJV ZKHQ WKH\ HPHUJH IURP WKH FRFRRQ WLRQ DQG VXUYLYDO 3DSDM   7R WHVW WKLV K\SRWKHVLV ZH FRP 6FKHGO   7KHUHIRUH IHPDOHV ZHUH NLOOHG GLUHFWO\ DIWHU SDUHG WKH UHVRXUFH DOORFDWLRQ SDWWHUQV RI VHYHUDO SLQH VDZIOLHV HPHUJHQFH E\ IUHH]LQJ DW ƒ& 7KH DPRXQW RI IDW ZDV GHWHU H[KLELWLQJ GLIIHUHQW RYLSRVLWLRQ VWUDWHJLHV ,Q SDUWLFXODU ZH FRQ PLQHG JUDYLPHWULFDOO\ IRU LQGLYLGXDO IHPDOHV IROORZLQJ WKH SUR VLGHUHG WKH LQYHVWPHQW RI IDW LQ HJJV YHUVXV VRPD DV IDW LV XVHG FHGXUH GHVFULEHG E\ (OOHUV   7ZR FRYHU VOLSV  î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ƒ&  GD\V IRU DOO VDZIO\ VSHFLHV DQG WKH GU\ ZHLJKW RI WKH VRPD 0$7(5,$/6 $1' 0(7+2'6
FRYHU VOLS  DQG WKH HJJ ORDG FRYHU VOLS   UHVSHFWLYHO\ ZDV 2ULJLQ RI LQVHFWV
GHWHUPLQHG 7KHQ  PO RI GLHWK\OHWKHU ZDV DGGHG WR HDFK FRYHU )LYH VSHFLHV RI 'LSULRQLGDH ZHUH VWXGLHG 7KH\ H[KLELW GLI VOLS DQG WKH IDW H[WUDFWHG IRU  K DW URRP WHPSHUDWXUH 7KHQ WKH IHUHQW RYLSRVLWLRQ VWUDWHJLHV DQG ODUYDO IHHGLQJ EHKDYLRXU GLHWK\OHWKHU ZDV FDUHIXOO\ FKDQJHG DQG WKH H[WUDFWLRQ FRQWLQXHG 0RUHRYHU WKH\ YDU\ LQ WKHLU RXWEUHDN WHQGHQF\ 7DEOH   7KH IRU DQRWKHU  K $IWHU WKLV WKH LQVHFW SDUWV ZHUH GULHG IRU  JUHJDULRXV IHHGLQJ 'LSULRQ SLQL / DQG 1HRGLSULRQ VHUWLIHU GD\V DW ƒ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ife history traits, outbreak tendency and origin of the laboratory cultures of pine sawfly species.
G. frutetorum Interme- Single eggs Fig. 1. Percentage of fat allocated to the egg load (mean ± SE) in females of five pine sawfly species. Significant differ- ences between means are indicated by different letters (P <0.05, t-tests on pairwise differences between parameter esti- still containing all the eggs), extracted for 24 h, 48 h or 96 h, so mates in the General Linearized Model).
we chose 48 h as a sufficient period for fat extraction.
In addition, the average egg size of each species was esti- tetorum invested about half of their fat in their eggs and (3) the mated by measuring the lengths and middle widths of 10 gregarious species, G. pallida, less than half of its fat (43%).
eggs/female and 5 females/species. The shape of the eggs of the Plasticity in the allocation of fat with increase in total fat was different species is similar (“banana”-like) and the volume (V) tested by the regression of the total fat content (x = total of an egg was calculated using V = S(width/2)² length.
fat/[unit BW]) on fat allocation (y = proportion of fat allocated Evaluation of data
to eggs) in a particular species. The basic allocation pattern remained unchanged in the gregarious species. However, in the 1. Number of eggs per female and average volume of eggs.
(semi-)solitary living D. similis and G. frutetorum the propor- 2.Total body weight (BW) [mg]: dry weight of soma + dry tion in the egg load decreased with increasing fat content (D. weight of egg load before fat extraction. BW was used as an similis: y = -1.299x + 0.824, F = 10.16, (1, 23), P < 0.005; G. frutetorum: y = -0.701x + 0.679, F = 3.34, (1, 51), P = 0.072).
3. Fat [mg]: absolute amount of fat in soma or egg load.
DISCUSSION
4. Fat content (Fat/[unit BW]): relative amount of fat in soma or egg load in relation to BW. Total fat/[unit BW] is the sum of The results of this study confirm that pine sawfly species differ in their pattern of fat allocation to eggs and soma. How- 5. Fat allocation: Proportion or percentage of total fat allocated ever, there is no clear relationship between the allocation pat- to the egg load or soma, respectively.
terns and the dichotomy in sawfly oviposition. D. pini and N.
sertifer
are gregarious and lay their eggs in one large cluster The statistical analysis was performed using linear regression (Pschorn-Walcher, 1982; Blümke & Anderbrant, 1997). Sur- and ANOVA by adopting General Linearized Models. Propor- vival of larvae increases with colony size (Lyons, 1962), tions were arcsine-transformed where appropriate. Means were probably because the feeding of the early instars is enhanced or compared by t-tests on pairwise differences between parameter large colonies are better able to defend themselves so reducing estimates in the General Linearized Model (Crawley, 1993).
the mortality risk per capita (Heitland & Pschorn-Walcher,1993; Codella & Raffa, 1995; Hunter, 2000). The data presented indicate that these sawfly species invested most of their fat in The sawfly species studied differed in several somatic and their egg load, producing more or larger eggs. D. pini had the reproductive traits (Table 2). The highest egg load was found in highest number of eggs (especially in comparison to the nearly D. pini, which was also the largest of the species (as indicated similar sized D. similis). N. sertifer had the least, but the largest by total BW). G. pallida and N. sertifer were significantly eggs of all the species. In contrast to the other European saw- smaller than the other species and had fewer eggs. However the flies, this species overwinters in the egg and presumably has to eggs of N. sertifer were larger than those of the other sawflies optimize its egg size in terms of energy reserves.
(Table 2: column 5). The sawflies also varied significantly in Distributing eggs singly or in several small clusters spatially their total fat content, with females of G. frutetorum accumu- can be seen as a risk-avoiding strategy, which is often associ- lating the most fat (Table 2: column 6). The fat content (fat/unit ated with cryptic colouration and behaviour of solitary feeding [BW]) of the egg load was significantly different from that of larvae (Prop, 1960). These species, in contrast to egg-clustering the soma in the gregarious species D. pini, N. sertifer and G. species, are likely to spend more time travelling and ovipositing, pallida, whereas the (semi-)solitary species D. similis and G. which are energetically costly. The results of this study lend to frutetorum invested equal amounts of fat in eggs and soma this contention, as the (semi-)solitary species G. frutetorum and D. similis allocated comparatively more fat to soma than D. pini The allocation of fat to the eggs differed between species and N. sertifer. Moreover, with increasing fat content, both (Fig. 1, ANOVA: F = 24.53, d.f.= (4, 207), P < 0.01). In princi- (semi-)solitary species increased their investment of fat in soma.
ple, three species groups can be distinguished: (1) the gregarious In addition, in the laboratory the longevity of the solitary G. fru- species N. sertifer and D. pini allocated most of their fat (about tetorum (10 days) was greater than that of the gregarious D. pini 60%) to their eggs, (2) the solitary species D. similis and G. fru- 7$%/(  $YHUDJH VRPDWLF DQG UHSURGXFWLYH WUDLWV RI QHZO\ HPHUJHG IHPDOHV RI ILYH SLQH VDZIO\ VSHFLHV 9DOXHV DUH PHDQV “ 6' Q QXPEHU RI IHPDOHV H[DPLQHG SHU VSHFLHV >X %:@ XQLW ERG\ ZHLJKW  0HDQV IROORZHG E\ GLIIHUHQW OHWWHUV DUH VLJQLILFDQWO\ GLI IHUHQW 3   HVWLPDWHG E\ WWHVWV RQ SDLUZLVH GLIIHUHQFHV EHWZHHQ SDUDPHWHU HVWLPDWHV LQ WKH *HQHUDO /LQHDUL]HG 0RGHO   6LJ QLILFDQW GLIIHUHQFHV EHWZHHQ WKH IDW FRQWHQW RI VRPD DQG HJJ ORDG IRU D SDUWLFXODU VDZIO\ VSHFLHV DW 3   DUH LQGLFDWHG E\ 6SHFLHV >PJ@
>PPñ@
>PJ@
>PJ@
 “ 
 “ D
 “ D
 “ 
 “ E
 “ E
 “ E
 “ E
 “ E
 “ F
 “ F
 “ F
 “ G
 “ F
 “ G
 “ 
 “ D
 “ G
 “ D
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|UNPDQ HW DO  WKDW DIIHFWV WKH VHOHFWLRQ RI SLQH VKRRWV )$,5 VSHFLILF 57' SURJUDPPH &7 µ3LQH VDZIO\ E\ IHPDOHV 8QIRUWXQDWHO\ QRWKLQJ LV NQRZQ DERXW WKH RYLSRVL SKHURPRQHV IRU VXVWDLQDEOH PDQDJHPHQW RI (XURSHDQ )RUHVWV WLRQ SUHIHUHQFH RI RWKHU SLQH VDZIOLHV DQG WKHUHIRUH LW LV GLIILFXOW 3+(52',3 ¶ ,W GRHV QRW QHFHVVDULO\ UHIOHFW WKH &RPPLVVLRQ¶V WR MXGJH KRZ IDVWLGLRXV * SDOOLGD LV LQ LWV FKRLFH RI RYLSRVLWLRQ YLHZ DQG LQ QR ZD\ DQWLFLSDWHV LWV IXWXUH SROLF\ LQ WKLV DUHD 6HYHUDO FRPSDUDWLYH VWXGLHV RI SHVW LQVHFWV DQG WKHLU QRQ 5()(5(1&(6
RXWEUHDN UHODWLYHV KDYH VXJJHVWHG DQ DVVRFLDWLRQ EHWZHHQ HJJ %((1$.(56 $07  &DUERK\GUDWH DQG IDW DV D IXHO IRU FOXVWHULQJJUHJDULRXVQHVV RI ODUYDH DQG RXWEUHDN IUHTXHQF\ LQVHFW IOLJKW $ FRPSDUDWLYH VWXG\ - ,QVHFW 3K\VLRO 
+DQVNL  :DOOQHU  /DUVVRQ HW DO   )RU WKH VSHFLHV FRQVLGHUHG LQ WKLV VWXG\ WKLV LV RQO\ SDUWLDOO\ WUXH 7KH $1'(5%5$17 2  2YLSRVLWLRQ SDWWHUQ DQG SDWWHUQ RI IDW DOORFDWLRQ WR VRPDWLF DQG UHSURGXFWLYH WLVVXHV EHKDYLRXU RI WKH SLQH VDZIO\ 1HRGLSULRQ VHUWLIHU +\PHQRS VHHPV WR EH D EHWWHU SUHGLFWRU RXWEUHDN WHQGHQF\ LQFUHDVHV ZLWK WHUD 'LSULRQLGDH  %XOO (QWRPRO 5HV  ±
DQ LQFUHDVH LQ WKH LQYHVWPHQW RI IDW LQ WKH HJJ ORDG ,Q WKH FDVH RI SLQH VDZIOLHV RXWEUHDNV PRVW OLNHO\ RFFXU ZKHQ SRSXODWLRQV SUHIHUHQFHV LQ SLQH VDZIOLHV D WUDGHRII EHWZHHQ ODUYDO DUH UHOHDVHG IURP WRSGRZQ FRQWURO +DQVNL 3DUYLDLQHQ  JURZWK DQG GHIHQFH DJDLQVW QDWXUDO HQHPLHV 2LNRV  ±
+DQVNL   :KHQ WKLV KDSSHQV VSHFLHV ZLWK D KLJK SRWHQWLDO IHFXQGLW\ PD\ H[KLELW D UDSLG SRSXODWLRQ EXLOGXS +DQVNL 5$))$ .)  &RQWULEXWLRQV RI IHPDOH RYL SRVLWLRQ SDWWHUQV DQG ODUYDO EHKDYLRU WR JURXS GHIHQFH LQ  :DOOQHU   ZKHUHDV VSHFLHV ZLWK D ORZ IHFXQGLW\ EXW FRQLIHU VDZIOLHV +\PHQRSWHUD 'LSULRQLGDH  2HFRORJLD 
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±
GXULQJ ODWHQW SHULRGV /DUVRQ 7HQRZ  6LPDQGO   7DNLQJ LQWR DFFRXQW WKH VPDOO VDPSOH VL]H ZH DQDO\VHG WKH 6&+(5,&+ (  'LH )RUVWLQVHNWHQ 0LWWHOHXURSDV %DQG 9 SURSRVHG UHODWLRQVKLS EHWZHHQ IDW DOORFDWLRQ DQG RXWEUHDN WHQ GHQF\ LQ SLQH VDZIOLHV VKRXOG EH UHJDUGHG DV D SUHOLPLQDU\ $16., ,  3LQH VDZIO\ SRSXODWLRQ G\QDPLFV SDWWHUQ SURF HVVHV SUREOHPV 2LNRV  ±
FRQFOXVLRQ )RU LQVWDQFH WKH RXWEUHDN WHQGHQF\ RI VSHFLHV VXFK DV * SDOOLGD FDQ YDU\ JHRJUDSKLFDOO\ 3VFKRUQ:DOFKHU   3$59,$,1(1 3  &RFRRQ SUHGDWLRQ E\ VPDOO 0RUHRYHU IDW DOORFDWLRQ PD\ EH LQIOXHQFHG E\ GLDSDXVH RU QX PDPPDOV DQG SLQH VDZIO\ SRSXODWLRQ G\QDPLFV 2LNRV 
WULWLRQ :H DUH DOVR DZDUH RI WKH IDFW WKDW RXU VWXG\ ZDV GRQH PDLQO\ RQ LQVHFWV UHDUHG LQ ODERUDWRU\ FXOWXUHV DQG ZH FDQQRW +(5= $  =XU (IIHNWLYLWlW XQG 6WUXNWXU GHU 3DUDVLWRLGHQJH H[FOXGH WKDW WKH VDZIOLHV ZHUH VXEMHFW WR DQ DUWLILFLDO VHOHFWLRQ PHLQVFKDIW YRQ 'LSULRQ SLQL / +\P 'LSULRQLGDH EHL ,GHDOO\ IDW DOORFDWLRQ SDWWHUQV DQG RWKHU OLIH KLVWRU\ WUDLWV RI ODWHQWHQ 'LFKWHQ GHV :LUWHV 3K' WKHVLV /XGZLJ ILHOG FROOHFWHG VSHFLPHQ IURP GLIIHUHQW JHRJUDSKLFDO RULJLQV DQG 0D[LPLOLDQV8QLYHUVLWlW 0QFKHQ  SS RI GLIIHUHQW SHVW VWDWXV VKRXOG EH FRPSDUHG WR GHWHUPLQH ZHWKHU :(,7=(/ &  2YLSRVLWLRQ GHWHUUHQFH E\ WKHUH LV D OLQN EHWZHHQ UHVRXUFH DOORFDWLRQ DQG SRSXODWLRQ FKHPLFDO VLJQDOV RI FRQVSHFLILF ODUYDH LQ 'LSULRQ SLQL +\PHQRSWHUD 'LSULRQLGDH DQG 3K\OORGHFWD YXOJDWLVVLPD &ROHRSWHUD &KU\VRPHOLGDH  (QWRPRO *HQ  ±
+817(5 $)  *UHJDULRXVQHVV DQG UHSHOOHQW GHIHQFHV LQ WKH 36&+251:$/&+(5 +  6\PSK\WD 'LSULRQLGDH ,Q VXUYLYDO RI SK\WRSKDJRXV LQVHFWV 2LNRV  ±
6FKZHQNH : HG  'LH )RUVWVFKl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±
PRO  ±
/<216 /$  7K H HIIHFW RI DJJUHJDWLRQ RQ HJJ DQG ODUYDO :$//1(5 :(  )DFWRUV DIIHFWLQJ LQVHFW SRSXODWLRQ VXUYLYDO LQ 1HRGLSULRQ VZDLQHL 0LGG +\PHQ 'LSULRQLGDH  G\QDPLFV GLIIHUHQFHV EHWZHHQ RXWEUHDN DQG QRQRXWEUHDN &DQ (QWRPRO  ±
VSHFLHV $QQX 5HY (QWRPRO  ±
3$3$- '5  2YDULDQ G\QDPLF DQG KRVW XVH $QQX 5HY :+((/(5 '  7KH UROH RI QRXULVKPHQW LQ RRJHQHVLV $QQX (QWRPRO  ±
5HY (QWRPRO  ± 
3523 1  3URWHFWLRQ DJDLQVW ELUGV DQG SDUDVLWHV LQ VRPH VSHFLHV RI WHQWKUHGLQLG ODUYDH $UFK 1pHUO =RRO 
5HFHLYHG -XQH   UHYLVHG 6HSWHPEHU   DFFHSWHG 1RYHPEHU  

Source: http://katz.entu.cas.cz/pdfarticles/7/eje_099_1_117_Herz.pdf

kiesels.info

Brain Formula ERHALTEN SIE IHRE GEISTIGEN FÄHIGKEITEN ◊ Wie verbessert Brain Formula meine geistige Gesundheit? Das letzte Jahrzehnt des 20. Jahrhunderts erlebte eine Explosion der Forschungim Bereich der Neurowissenschaften. Mittlerweile liegen fundierteAnhaltspunkte dazu vor, dass die speziellen Nährstoffe und die einzigartigenPflanzenextrakte von Brain Formula die Neuronen (

Microsoft word - creditor handbook

Pro Se Creditor’s Handbook: Information for Persons Owed Money by an Individual or Entity that has Filed for Bankruptcy Introduction This booklet is provided by the Office of the Clerk, United States Bankruptcy Court for the Eastern District of California. It has been prepared to respond to the questions frequently asked of Clerk's Office staff by non-lawyers. If you have a ques

Copyright 2014 Pdf Medic Finder