Douglas(fir+vulnerability+to+ cavita4on+and+adapta4on+to+ drought+events+ P.+Rozenberg,+G.+Dalla(Salda,+M.+Ruiz( Diaz,+A.S.+Sergent,+A.+Mar4nez(Meier,+ N.+Bréda,+L.+Sanchez,+J.C.+Bas4en+
• Drought(resistant+species+ • August+2003:+diebacks++ and+declines+ • Adapta4on+poten4al?+ Propor4on+of+observed+diebacks++
Propor4on+of+various++ species+in+the+2003+ diebacks+in+France+
Adapta4on+to+drought+events++ • Adapta4on+success:+ability+to+evolve++ • Evolu4onary+mechanism:+gene4c+varia4on+ and+heritability+of+adap4ve+traits+ – gene4c+varia4on+and+heritability:+large+number+of+ genotypes+and+individuals+ – adap4ve+trait:+related+with+fitness+(=+overall+ reproduc4ve+success)+ • Link+with+func4onal+processes:+resistance+to+cavita4on++ (intensive+and+4me(consuming+measurements(+ • ma4ng+ability,+fecundity+and/or+survivorship++
• Sample+collec4on+April+2006+ – Three+clones+ – Two+sites:+ • One+site+severely+affected+(C)+ • One+site+not+affected+(L)++
7575
C+site+ Chassenoix Lower+ring+density+ L+site+ Lartimache more+vulnerable+ +
• Measured+conduc4vity:+100+ 4mes+smaller+in+C+site+than+ in+L+site+ • In+C,+conduc4vity+clone+162:+ more+than+100+4mes+higher+ • Clone+162:+slower+growth,+ higher+wood+density,+more+ resistant+to+cavita4on+
circumference (cm)
70
6565 60
5555 50
4545
Higher+ring+density,+ more+resistant+ 0.5
162 1.0
1.5
1216 2.0
2.5
1407 3.0
3.5
clones Dalla(Salda,+G.,+A.+Mar4nez(Meier,+H.+Cochard,+et+P.+Rozenberg.+2009.+«+Varia4on+of+Wood+Density+and+ Hydraulic+Proper4es+of+Douglas(Fir+(+Pseudotsuga+Menziesii+(Mirb.)+Franco)+Clones+Related+to+a+Heat+ and+Drought+Wave+in+France.+»+Forest'Ecology'and'Management+257+(1).++
Microdensity+profiles+
Bark+
Pith+
Microdensity Profile
+
X(ray+picture+
2003+
1976+
+
Distance (X 25 microns)
5+
Conventional variables
0.8
0.9
Latewood density
0.6
0.7
Earlywood
Latewood Width
0.5
Earlywood Width
0.4
Ring Width
Earlywood density
0.3
Wood density (kg/dm3)
1.0
Latewood
0
20
40
60
Ring Width (X 10 microns)
Earlywood) (first)part))
80
• Sample+collec4on+March+2008+ – Seven+clones+X+three+trees+
• 2007+long+growing+season+at+Orléans++ – From+January+to+June,+warm+and+humid+ – From+July+to+September,+fresh+and+humid+ – October,+s4ll+warm+
P50 (MPa)
• +ring+2007+wide+and+conduc4ve:+wood+s4cks+ clone 1 2 3 4 5 6 7
-2.0 -2.0 -2.2 -2.2 -2.4 -2.4 -2.6 -2.6 -2.8 -2.8 -3.0 -3.0 0.20 0.22 0.20
0.26 0.24
0.24
0.28
0.30 0.32 0.28
Dalla(Salda,+G.,+A.+Mar4nez(Meier,+H.+Cochard,+et+P.+ Rozenberg.+2011.+«+Gene4c+Varia4on+of+Xylem+ Hydraulic+Proper4es+Shows+That+Wood+Density+Is+ Involved+in+Adapta4on+to+Drought+in+Douglas(Fir+ (Pseudotsuga'Menziesii'(Mirb.)).+»+Annals'of'Forest' Science+68+(4).++
0.32
Earlywood (first part) density (gr/cm3)
Hydraulic+proper4es+and+microdensity+variables+ P12) +++++++level+
+ clonal+
P88)
+ tree+
+ clonal+
+ tree+
+ clonal+
Mean+Ring+Density+
(0.56++
(0.76++
(0.70++
60.84))
Minimum+Density+
(0.57++
(0.76++
(0.71++
60.89))
Earlywood+Density+
(0.57+
(0.77+
(0.73+
60.85)
(0.71+ 60.85)
60.84)
60.97))
+
+ tree+
P50)
microdensity+
Earlywood)(first)part)) density) Maximum+Density+
(0.65+
60.85)
Latewood+Density+
(0.54+
(0.78+
Mortality:+dead+and+surviving+trees+ • 2004:+forêt+d’Orléans+(PhD+A.+Mar4nez(Meier)+ • From+the+90’s+to+2008+(succession+of+drought+ events)+(PhD+A.S.+Sergent)+
1990( 1991( 1999( 2003( 2005+
Causes+of+decline+ • Rela4onship+with+soil+water+ deficits+ Dieback(( Rate+ Non)calculé)
Rate+of+douglas(fir++ decline+
+ 8+ 6+ 4+ 2+ 0+
Risk'of'decline' threshold'
10
2001(
2003$ 2005& 2006+
2002( 2003$
2004& 2006+
+
0
Sergent,+Anne(Sophie,+Philippe+Rozenberg,+Nathalie+ Bréda+2012+«+Douglas(Fir+Is+Vulnerable+to+ Excep4onal+and+Recurrent+Drought+Episodes+and+ Recovers+Less+Well+on+Less+Fer4le+Sites+»,+Annals'of' Forest'Science,+1 12.++ +
+
50
+
100
+
150
+
200
+
250
Cumulated+soil+water+deficit++
Drought+events+quan4fied+by+soil+ water+balance+calcula4on,+Biljou©,+ INRA,+EEF+Nancy+ 10+
Rela4onship+with++ wood+density+
Mar4nez(Meier,+A.,+L.+Sanchez,+M.+ Pastorino,+L.+Gallo,+et+P.+ Rozenberg.+«+What+Is+Hot+in+Tree+ Rings?+the+Wood+Density+of+ Surviving+Douglas(firs+to+the+2003+ Drought+and+Heat+Wave.+»+Forest' Ecology'and'Management+256,+no+ 4+(2008).++
11+
0.20.30.40.50.60.70.80.9
Density+(kg.dm(3+
1990 1991 1992 1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003
Centre+ region+ 100
200
300
400
0.2 0.4 0.6 0.8
Density+(kg.dm(3+
0
Burgundy+ region+ 100
200
300
400
0.6
0.8
0
0.4
Midi(Pyrénées+ region+
0.2
Density+(kg.dm(3+
– Surviving trees – Dead trees
0
100
200
300
400
Length+of+microdensity+profile+(X+25+µm)+
Britez,+Manuela+Ruiz+Diaz,+Anne(Sophie+Sergent,+Alejandro+Mar4nez+Meier,+Nathalie+Bréda,+et+Philippe+ Rozenberg.+«+Wood+Density+Proxies+of+Adap4ve+Traits+Linked+with+Resistance+to+Drought+in+Douglas+Fir+ 12+ (Pseudotsuga+Menziesii+(Mirb.)+Franco)+».+Trees,+s.+d.,+1 16.++ +
Wood+density+variables+as+proxies+of:+
– Resistance+to+cavita4on+(=>+rela4onship+with+ func4onal+process)+ • • • •
Ring+density+ Minimum+density+ Earlywood+density+ Earlywood)(first)part))density)
– Survival+to+drought+event+(=>+rela4onship+with+fitness)+ • A+set+of+non(conven4onal+density+variables+ (BUT+relevant+variables+are+different+between+regions)+
• Poten4al+of+evolu4onary+adapta4on+within+the+ French+Douglas(fir+stands:+ – Gene4c+varia4on+ – Heritability+
Original variables
Conventional variables
Latewood Width
0.5
Earlywood Width
0.4
Ring Width
Earlywood density
0.6
0.8
Propor%on'of'' high'density'segment' MHD+ Density)threshold) Threshold++moving+ from+0.3+to+0.8+kg/m3+++++++++
0.4
0.6
0.7
Earlywood
MLD+ 0.2
Wood density (kg/dm3)
0.8
0.9
Latewood density
0.3
Wood density (kg/dm3)
1.0
1.0
Latewood
0
20
40
60
Ring Width (X 10 microns)
80
Low'density'segment' 0
50
100
Ring Width (X 25 microns)
Earlywood) (first)part))
150
Progeny+trials+ Three+provenances+X+18+progenies+ • Marblemount+ • Granite(Falls+ • Enumclaw+ Planted+in+three+progeny+trials+in+ France:+ • Orne,+Aude,+Vosges+
1600+increment+cores+
Proportion of High Density Segment Density threshold 0.4 (Survival) Narrow Sense Heritability 13
Genetic Variation
ORNE
2001 2004
2002
Site
Provenance Annual Rings Factors
11
1999 1998 2005
1999 1998
AUDE
2007 2003 2001
2002
8
GraniteFalls
VOSGES
10
2003 2005 2000
2000 Enumclaw
9
2004 2007
0.30
0.40
2008 2009 2006
VOSGES
0.20
mean of h2
Marblemount Enumclaw
ORNEMarblemount
12
2009 2006
mean of CV
0.50
2008
AUDE
GraniteFalls
Site
Provenance Annual Rings Factors
Earlywood (first part) density (Resistance to cavitation) Genetic Variation ORNE
10
ORNE
2006 2007 2005 2008 Enumclaw
1998
VOSGES
Site
8
2000 2002 1999
Provenance Annual
Enumclaw AUDE GraniteFalls
4
GraniteFalls
2003 2004 2005 2008 2007 2001
6
2003 2009 2004 2001
AUDE
2009 Marblemount
Marblemount
mean of CV
0.10 0.15 0.20 0.25 0.30 0.35 0.40 0.45
mean of h2
Narrow Sense Heritability
VOSGES
Site
2002 2000 1998 1999
Provenance Annual
Rings Factors
2006
Rings Factors
Provenance+trials:+signature+of+evolu4on+ Rela4onship+between+earlywood+ density+and+precipita4on+ + +
+
+
+
+
+
+
0.32 0.33 0.34 0.35 0.36 0.37 0.38 0.39
Earlywood+density++(+g/cm3)
+
R²+=+0.53+++(***)+
+
400
+ 800+ 1000+ 1200+ 1400+ 1600+ Annual+precipita4on+ in+the+natural+range+(mm)+ 600
Conclusion+ • Density+=+propor4on+of+cell+wall+ • For+several+density+variables+ – High+heritability+ – High+coefficient+of+varia4on+ – Signature+of+evolu4on+in+the+natural+range+
! High'potenAal'for'evoluAonary'adaptaAon'
Ques4ons+ • Rela4onships+between+survival+to+drought+and+ resistance+to+cavita4on?+ • Other+traits+for+resistance+to+soil+water+ deficit?+ • Different+density+variables:+ – Different+parts+of+the+ring?+ – Timing/intensity+of+soil+water+deficit?+
