(XU-(QWRPRO±
,661
(IIHFWVRIKRVWSODQWTXDOLW\RQRYHUZLQWHULQJVXFFHVVRIWKHOHDIEHHWOH
&KU\VRPHODODSSRQLFD&ROHRSWHUD&KU\VRPHOLGDH
(/(1$/=9(5(9$
6HFWLRQRI(FRORJ\'HSDUWPHQWRI%LRORJ\8QLYHUVLW\RI7XUNX7XUNX),1)LQODQGHPDLOHOH]YH#XWXIL
.H\ZRUGV&DUERQ&KU\VRPHODODSSRQLFD&ROHRSWHUD&KU\VRPHOLGDHGHOD\HGLQGXFLEOHUHVLVWDQFHKLEHUQDWLRQKRVWSODQW
TXDOLW\QLWURJHQSUHRYHUZLQWHULQJIHHGLQJ6DOL[P\UVLQLIROLDVXEVSERUHDOLVZHLJKWJDLQUDWHRYHUZLQWHULQJPRUWDOLW\
$EVWUDFW,LQYHVWLJDWHGHIIHFWVRIGHQVLW\GHSHQGHQWYDULDWLRQLQKRVWSODQWTXDOLW\RQDGXOWIHHGLQJDQGRYHUZLQWHULQJVXFFHVVRI
WKH OHDI EHHWOH &KU\VRPHOD ODSSRQLFD / 'XULQJ SUHRYHUZLQWHULQJ SHULRG DGXOW EHHWOHV ZHUH IHG LQ WKH ODERUDWRU\ RQ KRVW SODQWV
6DOL[ERUHDOLVRULJLQDWLQJIURPDDVLWHZLWKORZGHQVLW\SRSXODWLRQRI&ODSSRQLFDEDVLWHZLWKSHDNGHQVLW\RI&ODSSRQLFD
DQG F D SRVWRXWEUHDN VLWH %HHWOHV IHG RQ SODQWV IURP ORZ GHQVLW\ DQG SHDN GHQVLW\ VLWHV GHPRQVWUDWHG VLPLODU SHUIRUPDQFH
ZKHUHDVRQSODQWVIURPSRVWRXWEUHDNVLWHEHHWOHVIHGORQJHUJDLQHGPRUHZHLJKWDQGH[SHULHQFHGJUHDWHUPRUWDOLW\GXULQJRYHUZLQ
WHULQJ+LJKHUZDWHUFRQWHQWLQWKHVHEHHWOHVVXJJHVWVWKDWWKHPDLQUHDVRQIRULQFUHDVHGZLQWHUPRUWDOLW\LVLQVXIILFLHQWGHK\GUDWLRQ
RIWKHEHHWOHV¶ERGLHVSUHVXPDEO\GXHWRORZTXDOLW\IRRG7KHWRWDOFDUERQFRQFHQWUDWLRQLQKRVWSODQWIROLDJHH[SODLQHGYDULDWLRQLQ
PRVWEHHWOHSHUIRUPDQFHLQGLFHVZKLOHWRWDOQLWURJHQGLGQRWFRUUHODWHZLWKEHHWOHSHUIRUPDQFH'DPDJHLQGXFHGGHFUHDVHLQTXDOLW\
RIKRVWSODQWVIURPSRVWRXWEUHDNVLWHVGHOD\HGLQGXFLEOHUHVLVWDQFH',5DVVRFLDWHGZLWKLQFUHDVHLQFDUERQFRQWDLQLQJGHIHQVLYH
FRPSRXQGVZDVHDUOLHUIRXQGWRLQFUHDVHODUYDOPRUWDOLW\DQGWKXVFRQWULEXWHWRWKHGHFOLQHLQSRSXODWLRQGHQVLW\RI&ODSSRQLFD
5HVXOWVRIWKHFXUUHQWVWXG\VXJJHVWWKDW',5FDQDOVRGLVWXUESUHRYHUZLQWHULQJIHHGLQJDQGWKXVLQFUHDVHZLQWHUPRUWDOLW\RIDGXOW
EHHWOHVHQKDQFLQJSRVWRXWEUHDNGHQVLW\GHFOLQH
,1752'8&7,21
1DWXUDO PRUWDOLW\ RI LQVHFWV GXULQJ RYHUZLQWHULQJ PD\
EH YHU\ KLJK HVSHFLDOO\ LQ QRUWKHUQ DUHDV 'DQNV +RNNDQHQ /HDWKHU HW DO UHVXOWLQJ LQ VLJ
QLILFDQW HIIHFWV RQ LQVHFW SRSXODWLRQ G\QDPLFV *UHHQ
6XOOLYDQ :DOODFH 7HQRZ 1LOVVHQ
6XUYLYDO GXULQJ RYHUZLQWHULQJ GHSHQGV RQ ERWK
HQYLURQPHQWDO FKDUDFWHULVWLFV DQG LQWHUQDO SK\VLRORJLFDO
SURFHVVHV/HDWKHUHWDO+DQ%DXFH2QH
RI WKH PDMRU IDFWRUV ZKLFK GHWHUPLQHV WKH SHUIRUPDQFH
LQWHUPVRIVXUYLYDOZHLJKWGHYHORSPHQWGXUDWLRQDQG
SK\VLRORJ\ RI KHUELYRURXV LQVHFWV LV WKH TXDOLW\ RI WKHLU
KRVW SODQWV 6FULEHU 6ODQVN\ 0LOOHU 0LOOHU
.DUEDQ%DOGZLQ
&KDQJHVLQKRVWSODQWTXDOLW\IRUKHUELYRUHVZKLFKDUH
LQGXFHG E\ SODQW GDPDJH IURP KHUELYRU\ KDYH UHFHQWO\
UHFHLYHGPXFKDWWHQWLRQUHYLHZHGE\.DUEDQ%DOGZLQ
7KH GHFUHDVH LQ KRVW SODQWTXDOLW\ UHVXOWLQJ IURP
VHYHUHGDPDJHGXULQJSUHYLRXV\HDUVLVFDOOHGGHOD\HG
LQGXFLEOH UHVLVWDQFH ',5 $OWKRXJK ',5 DGYHUVHO\
DIIHFWV PDQ\ FKDUDFWHULVWLFV RI KHUELYRUH SHUIRUPDQFH
UHYLHZHG E\ +DXNLRMD 1HXYRQHQ .DUEDQ %DOGZLQ ',5 LPSDFW RQ RYHUZLQWHULQJ VXFFHVV
KDVQRWEHHQVWXGLHG',5LVDQLPSRUWDQWIDFWRULQLQVHFW
SRSXODWLRQG\QDPLFVEHFDXVH',5LVDGHQVLW\GHSHQGHQW
IXQFWLRQ RI KHUELYRUH GDPDJH L H KHUELYRUH SRSXODWLRQ
VL]HLWFRXOGSURYLGHWKHQHJDWLYHIHHGEDFNQHFHVVDU\WR
UHJXODWH IOXFWXDWLRQV LQ KHUELYRUH SRSXODWLRQV %HQ]
+DXNLRMD8QGHUZRRG
3RSXODWLRQV RI WKH ZLOORZIHHGLQJ OHDI EHHWOH &KU\
VRPHOD ODSSRQLFD / GHPRQVWUDWH SURQRXQFHG GHQVLW\
IOXFWXDWLRQV=YHUHYDHWDOD,QYHVWLJDWLRQRIIDF
WRUVDIIHFWLQJSRSXODWLRQG\QDPLFVRIWKLVVSHFLHVVKRZHG
WKDW QDWXUDO HQHPLHV SOD\ D PLQRU UROH LQ UHJXODWLRQ RI
GHQVLW\ G\QDPLFV =YHUHYD .R]ORY ZKLOH ',5
PD\ VLJQLILFDQWO\ FRQWULEXWH WR GHQVLW\ GHFUHDVH DIWHU DQ
RXWEUHDN =YHUHYD HW DO &KDQJHV LQ KRVW SODQW
TXDOLW\ LQGXFHG E\ SUHYLRXV GDPDJH SDUWLFXODUO\ KHUEL
YRUH RXWEUHDN UHVXOWHG LQ UHGXFHG IHFXQGLW\ DQG SUH
DGXOW VXUYLYDO LQFUHDVHG GXUDWLRQ RI GHYHORSPHQW DQG
GLVWXUEHGIHHGLQJEHKDYLRXURI&ODSSRQLFD=YHUHYD
.R]ORY =YHUHYD HW DO 7KHUHIRUH ,
K\SRWKHVL]HGWKDWGHFUHDVHLQKRVWSODQWTXDOLW\LH',5
PD\ LQFUHDVH ZLQWHU PRUWDOLW\ RI EHHWOHV GXH WR GLVWXU
EDQFH RI SUHRYHUZLQWHULQJ IHHGLQJ , WHVWHG WKLV
K\SRWKHVLVE\XVLQJKRVWSODQWVIURPVLWHVZKHUH&ODS
SRQLFD SRSXODWLRQV ZHUH RQ GLIIHUHQW VWDJHV RI GHQVLW\
IOXFWXDWLRQ DVVXPLQJ WKDW LQ SRVWRXWEUHDN VLWHV SODQWV
ZHUH RI ORZ TXDOLW\ GXH WR ',5 %HHWOH SHUIRUPDQFH
GXULQJWKHSHULRGEHWZHHQKDWFKLQJRIVXPPHUJHQHUDWLRQ
DQGWKHEHJLQQLQJRIGLDSDXVHDVZHOODVZLQWHUPRUWDOLW\
ZHUH VWXGLHG LQ & ODSSRQLFD EHHWOHV IHHGLQJ RQ SODQWV
IURPWKHWKUHHVLWHV
0$7(5,$/6$1'0(7+2'6
6WXG\DQLPDOV&KU\VRPHODODSSRQLFDDPHGLXPVL]HG±
PPOHQJWKOHDIEHHWOHLQ1:5XVVLDFOHDUO\SUHIHUV6DOL[ERUH
DOLV)ULHV1DVDUZKLFKLVFRPPRQLQWKHVWXG\DUHD=YHUHYD
HW DO E $GXOW EHHWOHV KLEHUQDWH LQ VRLO DQG VWDUW IHHGLQJ
DQG FRSXODWLQJ DW WKH WLPH RI OHDI IOXVK ZKLFK LQ RXU VWXG\
UHJLRQRFFXUVDWDERXWPLG-XQH/DUYDHIHHGRQZLOORZIROLDJH
IURPHDUO\-XO\WRHDUO\$XJXVWDQGSXSDWHRQOHDYHV%HHWOHVRI
WKH VXPPHU JHQHUDWLRQ HPHUJH LQ PLG$XJXVW IHHG IRU WZR
WKUHHZHHNVDQGWKHQGLJLQWRWKHVRLOIRURYHUZLQWHULQJ2YHU
ZLQWHULQJELRORJ\RIWKLVVSHFLHVLVXQNQRZQ
7$%/((IIHFWVRIKRVWSODQWLQGLYLGXDO%XVKDQGVLWHRIKRVWSODQWRULJLQ6LWHRQFKDUDFWHULVWLFVRI&KU\VRPHODODSSRQLFDEHIRUHDQG
GXULQJRYHUZLQWHULQJ*HQHUDO/LQHDU0RGHOWKH7\SH,,,6XPRI6TXDUHV
&KDUDFWHU
6RXUFH
GI
0HDQVTXDUH
)
3
)UHVKZHLJKWDWWKHEHJLQQLQJRIGLDSDXVH
6LWH
%XVK6LWH
(UURU
)UHVKZHLJKWJDLQGXULQJSUHRYHUZLQWHULQJIHHGLQJ 6LWH
%XVK6LWH
(UURU
)UHVKZHLJKWJDLQUDWH
6LWH
%XVK6LWH
(UURU
6LWH
%XVK6LWH
(UURU
6LWH
%XVK6LWH
(UURU
6LWH
%XVK6LWH
(UURU
6LWH
%XVK6LWH
(UURU
6LWH
%XVK6LWH
(UURU
6LWH
(UURU
)UHVKZHLJKWORVVGXULQJRYHUZLQWHULQJ
)UHVKZHLJKWDIWHURYHUZLQWHULQJ
'U\ZHLJKWDIWHURYHUZLQWHULQJ
:DWHUFRQWHQWLQEHHWOHV
'XUDWLRQRISUHRYHUZLQWHULQJIHHGLQJ
0RUWDOLW\GXULQJRYHUZLQWHULQJ
6LQFHPRUWDOLW\ZDVFDOFXODWHGRQSODQWVSHFLILFOHYHOHIIHFWRILQGLYLGXDOSODQWZLWKLQVLWHFRXOGQRWEHFDOFXODWHG
3RSXODWLRQGHQVLWLHVRI&ODSSRQLFDLQWHQVWXG\VLWHVORFDWHG
IURP NP 1( WR NP 6 RI WKH FLW\ RI 0RQFKHJRUVN .ROD
3HQLQVXOD 1: 5XVVLD Ħ 1 Ħ ( ZHUH UHFRUGHG LQ
PLG-XQHLQ E\PLQFRXQWV RIRYHUZLQWHUHGEHH
WOHVWKUHHFRXQWVSHUVLWHIRUGHWDLOVVHH=YHUHYDHWDOD
E ,Q VRPH VLWHV GHQVLWLHV UHPDLQHG QHDUO\ FRQVWDQW GXULQJ WKH
HQWLUHPRQLWRULQJSHULRGZKHUHDVLQRWKHUVLWHV,REVHUYHGVLJ
QLILFDQW GHQVLW\ IOXFWXDWLRQV XS WR IROG FKDQJH ZLWK WKH
SHDN YDOXHV DWWDLQHG DW GLIIHUHQW \HDUV =YHUHYD HW DO D
=YHUHYD.R]ORY
6WXG\VLWHVDQGKRVWSODQWV7RVWXG\WKHHIIHFWVRIGDPDJH
LQGXFHG FKDQJHV LQ KRVW SODQW TXDOLW\ RQ RYHUZLQWHULQJ RI &
ODSSRQLFD , IHG EHHWOHV LQ WKH ODERUDWRU\ RQ WKH IROLDJH RI LQGLYLGXDOVRI6ERUHDOLVIURPWKUHHVWXG\VLWHVILYHSODQWVSHU
VLWH7KHVHVLWHVGLIIHUHGLQFKDUDFWHULVWLFVRIOHDIEHHWOHSRSXOD
WLRQV L D VLWH ZLWK QRQIOXFWXDWLQJ ORZGHQVLW\ SRSXODWLRQ
ZKLFK VKRZHG QR VLJQLILFDQW GHQVLW\ FKDQJHV VLQFH ORZ
GHQVLW\VLWHKHUHDIWHULLDVLWHZLWKIOXFWXDWLQJSRSXODWLRQRQ
WKHVWDJHRIGHQVLW\LQFUHDVHRUSHDNGHQVLW\SHDNGHQVLW\VLWH
LLLDIOXFWXDWLQJSRSXODWLRQRQWKHVWDJHRISRVWRXWEUHDNGHQ
VLW\GHFOLQHSRVWRXWEUHDNVLWH
([SHULPHQWDOGHVLJQ$ERXWODVWLQVWDUODUYDHRI&ODS
SRQLFDFROOHFWHGRQ-XO\LQWKHSHDNGHQVLW\VLWHZHUH
IHG LQ WKH ODERUDWRU\ RQ IROLDJH RI 6 ERUHODLV IURP WKH ORZ
GHQVLW\VLWHLHQRWGDPDJHGGXULQJWKHFXUUHQWRULQWKHSUH
YLRXV\HDUVXQWLOSXSDWLRQ$IWHUKDWFKLQJZKLFKRFFXUUHG-XO\
± HDFK EHHWOH ZDV SODFHG LQ D PO YLDO DQG UDQGRPO\
DOORFDWHGWRRQHRIWKHVWXG\SODQWV±EHHWOHVSHUSODQW
LQ WRWDO )UHVK ZLOORZ OHDYHV ZHUH DGGHG WR YLDOV HYHU\
GD\ DQG WKH GD\ ZKHQ LQMXULHV RQ WKH IROLDJH ZHUH QRW IRXQG
ZDV FRQVLGHUHG DV WKH GD\ RI IHHGLQJ FHVVDWLRQ DQG WKH EHJLQ
QLQJ RI GLDSDXVH 2Q $XJXVW IUHVK VSKDJQXP PRVV ZDV
DGGHGWRHDFKYLDODQGYLDOVZHUHSODFHGIRURYHUZLQWHULQJLQWR
D FHOODU ZKHUH WKH WHPSHUDWXUH GLG QRW IDOO EHORZ ±ƒ& ,Q
VSULQJ0D\EHHWOHVZHUHUHPRYHGIURPWKHFHOODUDQGHDFK
EHHWOHZDVFODVVLILHGDVGHDGRUDOLYH
(DFK EHHWOH ZDV ZHLJKHG L LPPHGLDWHO\ DIWHU KDWFKLQJ DQG
KDUGHQLQJEHIRUHWKHVWDUWRIIHHGLQJ:+ZHLJKWDWKDWFKLQJ
LL WKH QH[W GD\ DIWHU EHHWOHV VWRSSHG IHHGLQJ DQG HPSWLHG WKH
JXW :' ZHLJKW DW WKH EHJLQQLQJ RI GLDSDXVH DQG LLL DIWHU
RYHUZLQWHULQJ:2EHIRUHEHHWOHVVWDUWHGIHHGLQJ0D\
VXUYLYRUV RQO\ 'U\ ZHLJKW RI WKH EHHWOHV WKDW VXUYLYHG DIWHU
RYHUZLQWHULQJ ': ZDV GHWHUPLQHG DIWHU GU\LQJ DW ƒ& WR
FRQVWDQW ZHLJKW LQ DLU 'XUDWLRQ RI SUHRYHUZLQWHULQJ IHHGLQJ
ZDV UHFRUGHG ' 6HYHUDO SDUDPHWHUV ZHUH FDOFXODWHG ZHLJKW
JDLQ GXULQJ SUHRYHUZLQWHULQJ IHHGLQJ :* :'±:+:+
ZHLJKW JDLQ UDWH GXULQJ SUHRYHUZLQWHULQJ IHHGLQJ :*5
:*' ZHLJKW ORVV GXULQJ RYHUZLQWHULQJ :/ :2±
:':'DQGERG\ZDWHUFRQWHQW :2±'::2
&DUERQDQGQLWURJHQDQDO\VLV$VDPSOHRIOHDYHVZDVFRO
OHFWHG IURP HDFK RI LQGLYLGXDOV RI 6 ERUHDOLV OHDYHV ZHUH
WDNHQIURPWKHPLGGOHSDUWRIWKHVKRRWVWRDYRLGDIIHFWVRIOHDI
DJH/HDYHVZHUHGULHGDWƒ&DQGWKHQJURXQG3ODQWVSHFLILF
WRWDO FDUERQ DQG QLWURJHQ ZHUH DQDO\]HG E\ /(&2 &+1
0DFURVDPSOH (OHPHQWDO $QDO\VHU /(&2 &RUSRUDWLRQ 6W
-RVHSK0LFKLJDQ
VARIATE procedure and variables not fitting normality
assumption were ln(x+1) transformed. Correlations between
beetle characteristics and the total carbon and nitrogen content
of the host plant were calculated on plant-specific level by Pearson’s correlation coefficient (SAS, 1990).
RESULTS
Both the site of host-plant origin and plant individual
affected beetle weight at the beginning of diapause, as
well as weight gain and weight gain rate during preoverwintering feeding (Table 1). Weight loss varied with
plant individual, but was not explained by the site of plant
origin. In contrast, duration of pre-overwintering feeding
depended on host-plant site, but did not depend on
among-plant variations within a site. The site of host
plant origin affected winter mortality (Table 1). Beetles
that fed on plants from the post-outbreak site gained more
weight compared to beetles fed on plants from lowdensity or peak-density sites (Fig. 1a, c). Higher weight
was attained due to both longer period of feeding and
higher weight gain rate (Fig. 1b, g). Weight loss in beetles that survived during overwintering was similar in all
study sites (Fig. 1d). Therefore, surviving beetles fed on
post-outbreak plants had higher weight after overwintering (Fig. 1e), mostly due to higher water content in
their bodies (Fig. 1f), whereas dry weight of the beetles
did not depend on host plant quality (Table 1). Beetles
which fed on plants from post-outbreak site suffered
higher winter mortality (Fig. 1h).
Although total carbon content in foliage of S. borealis
was similar in the three study sites (F2, 12 = 0.30, P =
0.75), weight gain, weight gain rate, weight at the beginning of overwintering, and duration of pre-overwintering
feeding positively correlated with plant-specific concentrations of carbon (Fig 2a, c, e, g). In contrast, total plant
nitrogen varied among study sites (F2, 12 = 19.9, P =
0.0002), but did not explain any of the studied beetle
characteristics (Fig. 2b, d, f, h).
DISCUSSION AND CONCLUSIONS
Fig. 1. Effect of the site of host-plant origin on performance
indices of Chrysomela lapponica. a – beetle fresh weight gain; b
– weight gain rate during pre-overwintering feeding; c – beetle
fresh weight at the beginning of overwintering; d – weight loss
during overwintering; e – fresh weight after overwintering; f –
body water content; g – duration of pre-overwintering feeding;
h – beetle mortality during overwintering. Sites differ in the
stage of C. lapponica population density fluctuation: postoutbreak site, peak-density site and low-density site. Each mean
is based on five plant-specific values; bars indicate standard
errors. Bars marked with different letters differ from each other
at the probability level P < 0.05 (Duncan’s multiple range test).
Data analysis. Effects of host-plant individual and of the site
of plant origin on leaf beetle characteristics were analyzed using
the General Linear Model (type III sum of squares); site-specific
means were compared with Duncan’s multiple range test. Beetle
mortality during overwintering was assessed at the plantspecific level, and therefore only the effect of site could be
assessed. Normality of all variables was checked by SAS UNI-
Effect of pre-overwintering feeding on winter
mortality
Overwintering mortality in insects may result from both
inadequate cold hardiness preparation and exhaustion of
energy reserves, which both can be significantly influenced by pre-winter conditions (Leather et al. 1993; Han
& Bauce, 1998). Individuals that accumulate larger
amounts of nutritive reserves during pre-overwintering
feeding have usually higher probability to survive during
the overwintering. For example, heavy individuals of
Meligethes aeneus overwintered successfully, whereas
low-weight beetles did not survive (Hokkanen, 1993). In
contrast, C. lapponica beetles, which fed on plants from
post-outbreak site and gained higher wet weight, showed
larger winter mortality. Comparison of dry weight
allowed the explanation of the high weight of these beetles by higher water content in beetle body.
Body water plays an important role in arthropod cold
hardiness: partial dehydration, which in many arthropods
is a necessary part of preparation for overwintering (Salt,
191
Fig. 2. Fresh beetle weight at the beginning of overwintering (a, b), pre-overwintering fresh weight gain (c, d); weight gain rate (e,
f); and duration of pre-overwintering feeding (g, h) of Chrysomela lapponica in relation to total carbon (a, c, e, g) and total nitrogen
(b, d, f, h) in Salix borealis foliage. Pearson’s correlation coefficient is calculated on the basis of 15 plants-specific values.
1956; van der Laak, 1982; Ring, 1982; Block, 1996;
Danks, 2000), may significantly affect cold resistance in
both freezing tolerant and freezing susceptible forms
(Block, 1996) by increasing cryoprotectant concentrations
and reducing ice nucleator activity (Gehrken, 1985;
Rickards et al., 1987; Pugh, 1994; Lee et al., 1996).
Respectively, high body water content negatively affects
cold hardiness and winter survival in insects (Vengorek,
1958; Cannon et al., 1985). Therefore it is likely that high
water content in high-weight beetles of C. lapponica may
be the main reason of increased winter mortality in popu192
lations feeding on plants from post-outbreak sites. It may
be suggested that individuals, which died during overwintering, had even higher water content than the survivors,
and this excess of water caused their death.
Many factors influence the body water content of
insects; in particular, feeding activity (Block, 1996) and
food quality (House, 1969; Murugan & George, 1992).
For example, phytophagous larvae of the spurge hawk
moth Celerio euphorbiae (L.) had higher water content
when fed on unbalanced artificial diet, whereas proportion of water in the diet had no effect on body water con-
WHQWLQODUYDH+RXVH%RG\ZDWHUFRQWHQWLQ&ROR
UDGR SRWDWR EHHWOH DW WKH EHJLQQLQJ RI GLDSDXVH ZDV
KLJKHULQEHHWOHVZLWKDORQJHUSUHRYHUZLQWHULQJIHHGLQJ
SHULRGDQGWKHVHEHHWOHVVXIIHUHGKLJKHUZLQWHUPRUWDOLW\
.RYDOVND 8VKDWLQVND\D +RZHYHU WR WKH
EHVWRIP\NQRZOHGJHDOPRVWQRWKLQJLVNQRZQDERXWWKH
OLQNVEHWZHHQKRVWSODQWFKDUDFWHULVWLFVDQGRYHUZLQWHULQJ
VXFFHVVRIKHUELYRUHV
3ODQWFKDUDFWHULVWLFVDIIHFWLQJSUHRYHUZLQWHULQJ
IHHGLQJ
(DUOLHUVWXGLHVGHPRQVWUDWHGWKDWLQGLYLGXDOVRI6ERUH
DOLVJURZLQJ LQSRVWRXWEUHDN VLWHVDUHSRRU TXDOLW\IRRG
IRU ERWK ODUYDH DQG DGXOWV RI & ODSSRQLFD 2Q WKHVH
SODQWV ODUYDH GHPRQVWUDWHG KLJKHU PRUWDOLW\ DQG ORZHU
FRQVXPSWLRQ UDWHV ZKLFK UHVXOWHG LQ SURORQJHG GHYHORS
PHQW DQG ORZHU SXSDO ZHLJKW =YHUHYD HW DO DGXOWVDOVRH[KLELWHGGLVWXUEHGIHHGLQJEHKDYLRU=YHUHYD
.R]ORY =YHUHYDHWDO $OOWKHVHVWXGLHV
GHPRQVWUDWHG WKDW GHFUHDVH LQ KRVWSODQW TXDOLW\ ZDV
FDXVHG E\ KHUELYRUH GDPDJH GXULQJ RXWEUHDN LH GDP
DJHG SODQWV KDYH GHYHORSHG GHOD\HG LQGXFLEOH UHVLVWDQFH
',5 7KH GHILQLWLRQ RI ',5 LV EDVHG RQ WKH HIIHFWV
REVHUYHG LQ KHUELYRUHV DQG FRQWDLQV QR DVVXPSWLRQV
DERXW SODQW FKDUDFWHULVWLFV UHVSRQGLQJ WR GDPDJH
+DXNLRMD .DUEDQ %DOGZLQ $OWHUDWLRQV
LQ SODQW FKHPLVWU\ PRVW IUHTXHQWO\ XQGHUO\ ',5 .DUEDQ
%DOGZLQEXWSK\VLFDOSODQWFKDUDFWHULVWLFVVXFK
DVVSHFLILFOHDIZHLJKWWKRXJKQHVVILEUHFRQWHQWSXEHV
FHQFH VSLQLQHVV PD\ DOVR FKDQJH LQ UHVSRQVH WR KHUEL
YRU\%DXUHWDO7DOODP\5DXSS7UXPEOH
HWDO,Q6ERUHDOLVWKHORZTXDOLW\RISUHYLRXVO\
GDPDJHG SODQWV ',5 ZDV ODUJHO\ H[SODLQHG E\ FKDQJHV
LQOHDISK\VLFDOWUDLWVLQFOXGLQJLQFUHDVHLQOHDIWKLFNQHVV
WRXJKQHVV KDLULQHVV DQG SURSRUWLRQ RI XQFRQVXPDEOH
EXONLQOHDIWLVVXHV=YHUHYDHWDO=YHUHYDXQSXE
OLVKHG 7KHVH FKDQJHV PD\ FDXVH QXWULHQW GHILFLHQFLHV
UHODWHG WR DQ LQFUHDVHG SURSRUWLRQ RI ILEHU $JDUZDOO
DQGFDQUHGXFHIRRGLQWDNH&KU\VRPHODODSSRQLFD
EHHWOHV SUHSDULQJ IRU RYHUZLQWHULQJ VHHP WR EH WU\LQJ WR
FRPSHQVDWH GHFUHDVHG IRRG LQWDNH RQ WRXJK DQG KDLU\
SODQWV IURP SRVWRXWEUHDN VLWH E\ ORQJHU IHHGLQJ +RZ
HYHU KLJKHU ZLQWHU PRUWDOLW\ LQGLFDWHV WKDW EHHWOHV IDLOHG
WRFRPSHQVDWHIRUWKHQXWULHQWGHILFLHQF\7KLVUHVXOWLVLQ
OLQHZLWKWKHGDWDRQODUYDOIHHGLQJDOWKRXJKODUYDHRI&
ODSSRQLFDIHGORQJHURQLQGXFHG6ERUHDOLVWKH\KDGVWLOO
JDLQHGORZHUZHLJKWDWSXSDWLRQDQGVXIIHUHGKLJKHUPRU
WDOLW\=YHUHYDHWDO
$PRQJSODQWYDULDWLRQLQZHLJKWJDLQDQGIHHGLQJGXUD
WLRQ RI & ODSSRQLFD ZHUH QRW H[SODLQHG E\ WRWDO IROLDU
QLWURJHQ ZKLFK RIWHQ VHUYHV DV D PHDVXUH RI KRVWSODQW
TXDOLW\ IRU KHUELYRUHV HJ 0DWWVRQ ,Q FRQWUDVW
WKH IHHGLQJ FKDUDFWHULVWLFV RI & ODSSRQLFD ZHUH FRUUH
ODWHG ZLWK WRWDO IROLDU FDUERQ +LJK FDUERQ FRQWHQW LQ D
SODQWLVWUDGLWLRQDOO\DWWULEXWHGWRLQFUHDVHGUHVRXUFHDOOR
FDWLRQWRFDUERQFRQWDLQLQJVHFRQGDU\FRPSRXQGVXVXDOO\
IXQFWLRQLQJ DV GHIHQVH DOVR LQGXFHG DJDLQVW KHUELYRUHV
&ROH\ HW DO +HUPV 0DWWVRQ .DUEDQ %DOGZLQ%XWLQ6ERUHDOLVLQGXFHGGHIHQFHVFDQ
XQOLNHO\ EH DWWULEXWHG WR DOOHORFKHPLFDOV =YHUHYD .R]ORY =YHUHYD 2ULDQV XQSXEOLVKHG GDWD
%HVLGHV KLJK FRQWHQW RI VRPH VHFRQGDU\ FKHPLFDOV
KLJKHUFDUERQLQOHDYHVPD\LQGLFDWHDQLQFUHDVHLQVWUXF
WXUDOFDUERK\GUDWHVLH³ILEUH´VXFKDVFHOOXORVHZKLFK
DUH FORVHO\ FRUUHODWHG WR OHDI WRXJKQHVV 6LQFH LQGXFHG
ZLOORZV DUH PRUH KDLU\ DQG WRXJK WKH KLJK FRQWHQW RI
VWUXFWXUDO FDUERK\GUDWHV LQ WKHVH SODQWV PD\ EH WKH PRVW
SODXVLEOHH[SODQDWLRQIRUVWURQJFRUUHODWLRQEHWZHHQSUH
RYHUZLQWHULQJ EHHWOH SHUIRUPDQFH DQG WRWDO IROLDU FDUERQ
IRXQG LQ & ODSSRQLFD $OWKRXJK WKH PHFKDQLVP OHDGLQJ
WR LQFUHDVHG ZDWHU FRQWHQW LQ EHHWOHV IHHGLQJ RQ WRXJK
SODQWV UHPDLQV XQNQRZQ LW PD\ EH UHODWHG WR LQFUHDVHG
IHHGLQJGXUDWLRQLQDWWHPSWVWRFRPSHQVDWHORZFRQVXPS
WLRQUDWHVDQGQXWULWLYHGHILFLHQF\
&RQVHTXHQFHVIRUSRSXODWLRQG\QDPLFV
$OWKRXJK ',5 LV FRQVLGHUHG DQ LPSRUWDQW IDFWRU UHJX
ODWLQJ KHUELYRUH SRSXODWLRQ G\QDPLFV 8QGHUZRRG
VRPH FDOFXODWLRQV VKRZ WKDW DGYHUVH HIIHFWV RI
',5 RQ SUHDGXOW LQVHFW SHUIRUPDQFH DUH QRW HQRXJK WR
FDXVHVLJQLILFDQWGHQVLW\GHFOLQH.DLWDQLHPLHWDO
(IIHFWRI',5RQVXUYLYDORIQRQIHHGLQJVWDJHVZDVRYHU
ORRNHG LQ HDUOLHU VWXGLHV DOWKRXJK IRRG TXDOLW\ LQ SUH
RYHUZLQWHULQJSHULRGPD\KDYHVWURQJLPSDFWRQVXUYLYDO
RIRYHUZLQWHULQJLQVHFWV$EVROXWHYDOXHVRIZLQWHUPRU
WDOLW\ZHUHUHODWLYHO\ORZLQP\H[SHULPHQWVZKHUHWHP
SHUDWXUHV GLG QRW GURS EHORZ ±ƒ& EXW GHWULPHQWDO
HIIHFWVRI',5RQRYHUZLQWHULQJVXUYLYDORI&ODSSRQLFD
PD\ EH PXFK PRUH GUDPDWLF LQ WKH QDWXUDO HQYLURQPHQW
ZKHUHPLQLPXPWHPSHUDWXUHVRIWKHXSSHUVRLOOD\HUPD\
UHDFK±ƒ&.R]ORY+DXNLRMD,QWKHVHFRQGL
WLRQVDQLQFUHDVHLQERG\ZDWHUFRQWHQWPD\KDYHDPRUH
VHYHUH LPSDFW RQ LQVHFW VXUYLYDO YLD FKDQJHV LQ VXSHU
FRROLQJFDSDFLW\WKDQLWZDVREVHUYHGLQWKHPLOGFRQGL
WLRQVRIP\H[SHULPHQW
%HVLGHV WKH GLUHFW HIIHFW RI ',5 RQ WKH PRUWDOLW\ RI
RYHUZLQWHULQJ EHHWOHV GXH WR SK\VLRORJLFDO UHDVRQV
LQFUHDVHGGXUDWLRQRISUHRYHUZLQWHULQJIHHGLQJLQQDWXUH
SURORQJV H[SRVXUH WR SUHGDWLRQ LQ DXWXPQ ZKHQ ELUGV
IHHGHVSHFLDOO\DFWLYHO\7KHUHVXOWVRIP\VWXG\LQGLFDWH
WKDW DQ LQFUHDVH LQ ZLQWHU PRUWDOLW\ FDXVHG E\ ',5 PD\
FRQWULEXWH VLJQLILFDQWO\ WR WRWDO JHQHUDWLRQ PRUWDOLW\ ,Q
FRPELQDWLRQ ZLWK RWKHU GHWULPHQWDO HIIHFWV RI ',5 VXFK
DVUHGXFWLRQLQIHFXQGLW\DQGODUYDOPRUWDOLW\DGXOWPRU
WDOLW\ GXULQJ RYHUZLQWHULQJ PD\ OHDG WR FRQVLGHUDEOH
SRSXODWLRQGHQVLW\GHFOLQHDIWHURXWEUHDN
$&.12:/('*(0(176 , WKDQN 'PLWU\ 1RYLNRY IRU DVVLV
WDQFHLQWKHODERUDWRU\ZRUN$UMD7HUYDKDXWDIRUWKHDQDO\VLVRI
FDUERQDQGQLWURJHQLQZLOORZOHDYHVDQG0.R]ORYIRUYDOX
DEOH FRPPHQWV WR DQ HDUOLHU GUDIW RI WKH PDQXVFULSW 7KH ZRUN
ZDVVXSSRUWHGE\0DMDQG7RU1HVVOLQJ)RXQGDWLRQ7XUNX8QL
YHUVLW\ )RXQGDWLRQ DQG 8QLYHUVLW\ RI -RHQVXX YLD JUDQW WR 3
1LHPHOl
5()(5(1&(6
$*$5:$// 5$ 0RUSKRORJLFDO FKDUDFWHULVWLFV RI VXJDU
FDQHDQGLQVHFWUHVLVWDQFH(QWRPRO([S$SSO±
%$85 5 %,1'(5 6 %(1= * 1RQJODQGXODU OHDI WUL
FKRPHV DV VKRUWWHUP LQGXFLEOH GHIHQFH RI WKH JUD\ DOGHU
$OQXVLQFDQD/DJDLQVWFKU\VRPHOLGEHHWOH$JHODVWLFDDOQL
/2HFRORJLD±
%(1= * 1HJDWLYH IHHGEDFN E\ FRPSHWLWLRQ IRU IRRG DQG
VSDFH DQG E\ F\FOLF LQGXFHG FKDQJHV LQ WKH QXWULWLRQDO EDVH
DV UHJXODWRU\ SULQFLSOHV LQ WKH SRSXODWLRQ G\QDPLFV RI WKH
ODUFKEXGZRUP =HLUDSKHUDGLQLDQD*XHQHH /HS7RUWULFL
GDH=$QJHZ(QWRPRO±
%/2&.:&ROGRUGUDXJKW±WKHOHVVHURIWZRHYLOVIRUWHU
UHVWULDODUWKURSRGV"(XU-(QWRPRO±
&$1121 5- %/2&. : &2//(77 *' /RVV RI VXSHU
FRROLQJ DELOLW\ LQ &U\SWRS\JXV DQWDUFWLFXV &ROOHPEROD ,VR
WRPLGDHDVVRFLDWHGZLWKZDWHUXSWDNH&U\R/HWWHUV±
&2/(<3'%5<$17-3&+$3,1)6,,,5HVRXUFHDYDLO
DELOLW\ DQG SODQW DQWLKHUELYRUH GHIHQVH 6FLHQFH ±
'$1.6+90RGHVRIVHDVRQDODGDSWDWLRQLQWKHLQVHFWV
,:LQWHUVXUYLYDO&DQ(QWRPRO±
'$1.6 +9 'HK\GUDWLRQ LQ GRUPDQW LQVHFWV - ,QVHFW
3K\VLRO±
*(+5.(1 8 3K\VLRORJ\ DQG GLDSDXVH RI WKH DGXOW EDUN
EHHWOH,SVDFXPLQDWXV*\OOVWXGLHGLQUHODWLRQWRFROGKDUGL
QHVV-,QVHFW3K\VLRO±
*5((1 *: /RZ ZLQWHU WHPSHUDWXUHV DQG WKH (XURSHDQ
SLQH VKRRW PRWK 5K\DFLRQLD EXROLDQD 6FKLII LQ 2QWDULR
&DQ(QWRPRO±
+$1 (1 %$86( ( 7LPLQJ RI GLDSDXVH LQLWLDWLRQ
PHWDEROLF FKDQJHV DQG RYHUZLQWHULQJ VXUYLYDO RI WKH VSUXFH
EXGZRUP &KRULVWRQHXUD IXPLIHUDQD (FRO (QWRPRO ±
+$8.,2-$ ( ,QGXFWLRQ RI GHIHQVHV LQ WUHHV $QQX 5HY
(QWRPRO±
+$8.,2-$ ( 1(8921(1 6 ,QVHFW SRSXODWLRQ G\QDPLFV
DQG LQGXFWLRQ RI SODQW UHVLVWDQFH 7KH WHVWLQJ RI K\SRWKHVHV
,Q %DUERVD 3 6FKXOW] -& HGV ,QVHFW 2XWEUHDNV $FD
GHPLF3UHVV6DQ'LHJR86$SS±
+(506 '$ 0$77621 :- 7KH GLOHPPDRI SODQWV WR
JURZRUWRGHIHQG45HY%LRO±
+2..$1(1 +07 2YHUZLQWHULQJ VXUYLYDO DQG VSULQJ
HPHUJHQFH LQ 0HOLJHWKHV DHQHXV HIIHFWV RI ERG\ ZHLJKW
FURZGLQJ DQG VRLO WUHDWPHQW ZLWK %HDXYHULD EDVVLDQD (QWR
PRO([S$SSO±
+286( +/ (IIHFWV RI GLIIHUHQW SURSRUWLRQV RI QXWULHQWV
RQLQVHFWV(QWRPRO([S$SSO±
.$,7$1,(0, 3 1(8921(1 6 1<<66g1(1 7 (IIHFWV RI
FXPXODWLYH GHIROLDWLRQV RQ JURZWK UHSURGXFWLRQ DQG LQVHFW
UHVLVWDQFHLQPRXQWDLQELUFK(FRORJ\±
.$5%$15%$/':,1-7,QGXFHG5HVSRQVHVWR+HUEL
YRU\ 7KH 8QLYHUVLW\ RI &KLFDJR 3UHVV &KLFDJR 86$ SS
.2=/2909+$8.,2-$(0LFURFOLPDWHFKDQJHVDORQJ
DVWURQJSROOXWLRQJUDGLHQWLQQRUWKHUQERUHDOIRUHVW]RQH,Q
8VR -/ %UHEELD &$ 3RZHU + HGV (FRV\VWHPV DQG
6XVWDLQDEOH 'HYHORSPHQW $GYDQFHV LQ (FRORJLFDO 6FLHQFHV
9RO &RPSXWDWLRQ 0HFKDQLFV 3XEOLFDWLRQV 6RXWKDPSWRQ
8.SS±
.29$/6.$ 7 (IIHFW RI SUHRYHUZLQWHULQJ SK\VLRORJLFDO
VWDWHRIWKH&RORUDGRSRWDWREHHWOH/HSWLQRWDUVDGHFHPOLQHDWD
6D\RQGLDSDXVHDQGPRUWDOLW\GXULQJRYHUZLQWHULQJ,Q3DY
ORYVN\(1HG&RORUDGR3RWDWR%HHWOHDQG0HWKRGVRILWV
&RQWURO ,VVXH $FDGHP\ RI 6FLHQFH RI WKH 8665 3XEO
+RXVH0RVFRZSS±>LQ5XVVLDQ@
/($7+(565:$/7(56.)$%$/(-67KH(FRORJ\
RI,QVHFW2YHUZLQWHULQJ&DPEULGJH8QLY3UHVV&DPEULGJH
SS
/(( 5( &267$1=2 -3 08*1$12 -$ 5HJXODWLRQ RI
VXSHUFRROLQJ DQG LFH QXFOHDWLRQ LQ LQVHFWV (XU - (QWRPRO
±
0$77621 :--U +HUELYRU\ LQ UHODWLRQ WR SODQW QLWURJHQ
FRQWHQW$QQX5HY(FRO6\VW±
0,//(5 -5 0,//(5 7$ HGV ,QVHFW3ODQW
,QWHUDFWLRQV6SULQJHU9HUODJ1HZ<RUNSS
0858*$1.*(25*($)HHGLQJDQGQXWULWLRQDOLQIOX
HQFH RQ JURZWK DQG UHSURGXFWLRQ RI 'DSKQLV QHULL /LQQ
/HSLGRSWHUD6SKLQJLGDH-,QVHFW3K\VLRO±
38*+ 3-$ 6XSHUFRROLQJ SRLQWV DQG ZDWHU FRQWHQW LQ
$FDUL$FWD2HFRO±
5,&.$5'6-.(//(1(50-6725(<.%6WUDWHJLHVRI
IUHH]H DYRLGDQFH LQ ODUYDH RI WKH JROGHQURG JDOO PRWK (SL
EOHPDVFXGGHULDQDZLQWHUSURILOHVRIDQDWXUDOSRSXODWLRQ-
,QVHFW3K\VLRO±
5,1*5$)UHH]LQJWROHUDQWLQVHFWVZLWKORZVXSHUFRROLQJ
SRLQWV&RPS%LRFKHP3K\VLRO$±
6$/75:,QIOXHQFHRIPRLVWXUHFRQWHQWDQGWHPSHUDWXUH
RQ FROGKDUGLQHVV RI KLEHUQDWLQJ LQVHFWV &DQ - =RRO ±
6$6,167,787(6$667$78VHU
V*XLGH9HUVLRQ6$6
,QVWLWXWH&DU\1RUWKHUQ&DUROLQD86$SS
6&5,%(5 -0 6/$16.< )-U 7KH QXWULWLRQDO HFRORJ\ RI
LPPDWXUHLQVHFWV$QQX5HY(QWRPRO±
68//,9$1 &5 :$//$&( '5 7KH SRWHQWLDO QRUWKHUQ
GLVSHUVDO RI WKH J\SV\ PRWK 3RUWKHWULD GLVSDU /HSLGRSWHUD
/\PDQWULLGDH&DQ(QWRPRO±
7$//$0<':5$8330-HGV3K\WRFKHPLFDO,QGXF
WLRQLQ+HUELYRUHV-:LOH\6RQV1HZ<RUNSS
7(12:2 1,/66(1$ (JJ FROGKDUGLQHVVDQG WRSRFOL
PDWLF OLPLWDWLRQV WR WKH RXWEUHDNV RI (SLUULWD DXWXPQDWD LQ
QRUWKHUQ)HQQRVFDQGLD-$SSO(FRO±
7580%/( -7 .2/2'1<+,56&+ '0 7,1* ,3 3ODQW
FRPSHQVDWLRQ IRU DUWKURSRG KHUELYRU\ $QQX 5HY (QWRPRO
±
81'(5:22' 1 7KH LQIOXHQFH RI LQGXFHG UHVLVWDQFH RQ
KHUELYRUHSRSXODWLRQV,Q$JUDYDO$$7X]XQ6%HQW(
HGV,QGXFHG3ODQW'HIHQVHVDJDLQVW3DWKRJHQVDQG+HUEL
YRUHV$363UHVV6W3DXO0LQQHVRWD86$SS±
86+$7,16.$<$56&RORUDGR3RWDWR%HHWOH/HSWLQRWDUVD
GHFHPOLQHDWD6D\3K\ORJHQ\0RUSKRORJ\3K\VLRORJ\(FRO
RJ\ $GDSWDWLRQ 1DWXUDO (QHPLHV ³1DXND´ 0RVFRZ SS>LQ5XVVLDQ@
9$1 '(5 /$$. 6 3K\VLRORJLFDO DGDSWDWLRQV WR ORZ WHP
SHUDWXUH LQ IUHH]LQJWROHUDQW 3K\OORGHFWD ODWLFROOLV EHHWOHV
&RPS%LRFKHP3K\VLRO$±
9(1*25(.9:ĉ*25(.:7KHVWXG\RIRYHUZLQWHULQJ
RI WKH&RORUDGR SRWDWREHHWOH RQWKHEDVH RILW¶V SK\VLRORJ\
,Q 3DYORYVN\ (1 HG &RORUDGR 3RWDWR %HHWOH DQG
0HWKRGV RI LWV &RQWURO ,VVXH $FDGHP\ RI 6FLHQFH RI WKH
86653XEO+RXVH0RVFRZSS±>LQ5XVVLDQ@
=9(5(9$ (/ .2=/29 09 $YRLGDQFH RI ZLOORZV
IURPPRGHUDWHO\SROOXWHGDUHDE\OHDIEHHWOH0HODVRPDODS
SRQLFD &ROHRSWHUD &KU\VRPHOLGDH HIIHFWV RI HPLVVLRQ RU
LQGXFHGUHVLVWDQFH"(QWRPRO([S$SSO±
=9(5(9$ (/ .2=/29 09 (IIHFWV RI SROOXWLRQ RQ
QDWXUDO HQHPLHV RI WKH OHDI EHHWOH 0HODVRPD ODSSRQLFD -
$SSO(FRO±
=9(5(9$(/.2=/29091(8921(16D3RSXODWLRQ
GHQVLW\DQGSHUIRUPDQFHRI0HODVRPDODSSRQLFD&ROHRSWHUD
&KU\VRPHOLGDHLQVXUURXQGLQJVRIVPHOWHUFRPSOH[(QYLURQ
(QWRPRO±
=9(5(9$(/.2=/29091(8921(16E'HFUHDVHLQ
IHHGLQJ QLFKH EUHDGWK RI 0HODVRPD ODSSRQLFD &ROHRSWHUD
&KU\VRPHOLGDH ZLWK LQFUHDVH LQ SROOXWLRQ 2HFRORJLD ±
=9(5(9$ (/ .2=/29 09 1,(0(/b 3 +$8.,2-$ ( 'HOD\HG LQGXFHG UHVLVWDQFH DQG LQFUHDVH LQ OHDI IOXFWXDWLQJ
DV\PPHWU\DVUHVSRQVHVRI6DOL[ERUHDOLVWRLQVHFWKHUELYRU\
2HFRORJLD±
=9(5(9$(/.2=/29091,(0(/b3(IIHFWVRIOHDI
SXEHVFHQFHLQ6DOL[ERUHDOLVRQKRVWSODQWFKRLFHDQGIHHGLQJ
EHKDYLRXU RI WKH OHDI EHHWOH 0HODVRPD ODSSRQLFD (QWRPRO
([S$SSO±
5HFHLYHG1RYHPEHUUHYLVHG-DQXDU\DFFHSWHG0DUFK