Douglas(fir+vulnerability+to+ cavita4on+and+adapta4on+to+ drought

Cumulated+soil+water+deficit++. Rate+of+douglas(fir++ decline+. Risk'of'decline' .... 2008. 2009. Site. Provenance Annual. Rings. Narrow Sense Heritability ...
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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?+