(¾#Y01ÊQË·
"#$%&'()*+,')ÌÍÎÏ(rÐbpqÑÒÓ[ÔXÕÖ#^_!

.#$!-{|)*×  Øpr~./012345(6789:;'YÙÚÛ?Ü?kQ(b­§Ý?><=>?@ABCDEFG8HIJKL[Þ#^_ß`àk#$XX(~n¶M !"N OPQIJRSTU

 'S|X#5cáâãVWX(Y01IZ[\ä"#ÈÙkQ(^_.å9æÚ(å]"#^_`a5$b5c(Y019æÚçèéê~n¶ëÙ01[IìX[d5)*'3$5([+,efg4(!hij[kl#$Y01m'¸

nopq+,rstuvw#$x m®ïðñòóôõk+,'  yzX5$ '{|}#( Hí¨îc34#$~n¶Iìö'W÷øù[ú~'}1}M  N_5Xr3(m£ûXXrQ(3þ­ösk#5c(SI["$XX(0n#Ý?>k#$b^x+,'(<?}1pq'!Q( 'æFä[d5~n¶  R¯n34#$^IJö
#IZ'(O[g5$HüZýjà

OX5~n¶ ms¡4#_¢£4#$R¯IJö[m[d#^_(SI'I35

501¤¥[¦§X(I[¦3X5$ ~¶  ¥¨©

£Þ34

nðñdX( ~×  ªF«=¬[Q­§®R¯°±.#$ k#nðñsIJö[dX
5(²³(´µ¶  m

5$_·D¸#^_(~_¹)*
º!(»~pq¼0(~½¾_¿À¦ÁÂ4#$5( 'Ã1¹Ä1(Å}0Ä1Æ¶Ç_X¾ÈÉ01k#$¾Æ!'[-.)-~n¶  R¯IJö[$ Ú[!¸#5c(m[d#^_-yOX5ôõ$X5<?

"% $&'MNæF[d5Iì(rÐ(O¹(À+,*í      (=FM7N !"#$%&'()

*&+,-./0&!1234&56785yz)2MnZ{e2|}~&S2&gP)

+M

6789:;<!=>?@ABC&)23QS&TQU

DEF)

GH;<!=IJ@AK

Rpdh

L%&MN-

.O23SPQ2 RS#&TQUP)23 x ¡¢c^£¤)2¥¦Q2VWXYZT[\]^_`abcZde2p>?!§¨©fUgP-

2ª«Q

.!123¬§&[1U­JM+Bp®

!

3 ))¬§

&[hRS#4i$jkKlYm&XY-

.O23M

nojpqrstuvwM
VWXY@A)UPQ
xURS#-

!123

./0&!1

23/0¯.dpd123±²ef&[°#Q

R&TQU³l. 

pⷫ᳓ᕈ䊎䊆䊦䊘䊥䊙䊷´µ¶· %,-*7

{A

X!"#$O+XdVdm%&'()(*+#,-. /  5LA&!1

2
0
 1
!2  / 0"1#2 / $"1" /  #  9:;jk2   / !
2  / "11
 234
1
"2 / 0 9:;jk&((B  8( && 4  ¡ '(51%&6/ !7   #%89:;)*+ &L¢#£b¤+¢#¥#<=>?

b¤¦c¤§(#¨  L@ABCDEFGHI#JK:MABN-

O#  >?

&©+A %&LAP
ª!"#  8#«¬ @ABCDEVWXYZ[\]^_`a`bc

FA#­®VWO+X#¯°[d#efg#hi& LG  %±²GQA9:;RS8TUB4O  9:;jk&l]AmRnop

4m³¨#´VWµ ¶-5qrstuGv4wx¹A(&ºc»{((·¸  #¼u^_`a`bc[d#½4  9:;#ef
¾¿[d#ÀÁ ÃO  9:;Rny8z{A|O

B5r(B4Â

(B4 9:;jk#l]& 9:;Rny8}~AVW¾¿[O#^_`a`bc[#ÄÅÆ  373WV#lcVWX?ÆÇÈÉÊË#47r    + #   3'(VW¾¿[##ÌÍÎ#4)*+ ᾏỈ㻞㻚㻡This work⍇⍈†Ἑᕝ※ὶᇦἙᕝ୰䞉ୗὶᇦ㞵Ỉ

G98/95Mo (‰)

㻞㻚㻜Archer and Vance (2008)㻝㻚㻡㻷㼍㼘㼕㼤㻌㻾㼕㼢㼑㼞㻺㼕㼘㼑㻌㻾㼕㼢㼑㼞㻌㻔㻹㼍㼕㼚㻕㻺㼕㼘㼑㻌㻾㼕㼢㼑㼞㻌㻔㼃㼔㼕㼠㼑㻕㻺㼕㼘㼑㻌㻾㼕㼢㼑㼞㻌㻔㻮㼘㼡㼑㻕㼂㼛㼘㼓㼍㻌㻾㼕㼢㼑㼞㻯㼔㼍㼚㼓㻌㻶㼕㼍㼚㼓㻌㻾㼕㼢㼑㼞㻻㼠㼠㼍㼣㼍㻌㻾㼕㼢㼑㼞㻯㼘㼑㼍㼞㻌㻯㼞㼑㼑㼗㻮㼞㼍㼔㼙㼍㼜㼡㼠㼞㼍㻭㼙㼍㼦㼛㼚㻝㻚㻜㻜㻚㻡ᆅẆᒾ▼㻜㻚㻜㻜㻚㻜㻜㻚㻡㻝㻚㻜㻝㻚㻡㻞㻚㻜䠄Error bar : r2 SD䠅

1/[Mo] (nmol-1 kg)VWO#.
µ ¶      !"#P!ÀÁÂÃ.ÄÅÆ=-P;DÇ+,-./

0Èz=¢£y/xyÉÊ/º»Ë(ÌY1234 5!"#2&64=¼½$%&'()*i=Í¨/ QRSÎÏ=fHz

7(89/:;(<1+,=>?

yI/ÌY=¼½@A@BCDEFGHCI

M=Ñ(ÒÓ44/ÔÕ!  =Q

BCKLMNOP

J=>?@A@QRSTUVi/(=BÐ

RSÖz×[P !

QØ

WXYZP4[\]^¢£y

_=`,a¨My/Ù!Ú.ZP=bcde(f40=g!/hi+,/

=j%¢!

5=!QRS)(zÛÜZ(&6k"#lmP5n4[/o2"#p&'(2q!

mÞß5¢xyÉÊCàá

5=rs2tP4 5!

{|.

=uv  =QRSÖ=°±DÝj=v/BÐM=ÑâãZ5¢(w,>?@A@BCN=VWw,xyz{$  äåQRSÖæDçè/$

|P4[(u}+,h~k/;!>?

QRSÝ

@A@BCNVWOPQRSTU(X  QRS¼½D×êZß5

0zh~Pz=

¢|4/ÀÀë34ì2y=

24[( 4/RS!B$=í¢£y/52ÁÂDîHkï£

={|.5234 5!
@B

Q Dé[2(45(x3J=  NZP5DP¬­!"#@B@B=e(!QRS $ 5y/OPQRS@B¬­!
%
&'()
!*(+Z @B/ ((+ ,"#   / !KLM

¡(VWZ/J2XYZ5u}M=+,=-%/

J¢=¬ ­')./(*, 
' ,*00=h~=+ 1/2,.+3%#n¢£5¢¤¥¦§(xy¢=QRS=¨M(©Pª«Z   - 4¬­1&5"#   P4¬­/®cd$=«¯°±²k5³´µZ¶y¬­/·

-$ 4
,


#    ¬­ ,-4 

¢zQ¸=°±¹4¬­"1!"  6, (6
&7"8#  ·=±!/ º»=      QRS¼½(¶P¾¿¢PP;/£ 