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The effect of ectomycorrhizal fungi on reducing the variation of seedling growth of Eucalyptus globulus
41
Agriculture, Ecosystems and Environment, 28 ( 1 9 8 9 ) Elsevier Science Publishers B.V., Amsterdam Printed in Czechoslovakia
41-46
THE EFFECT OF EC'TOMYCORRHIZAL FUNGI ON REDUCING THE VARIATION OF' SEEDLING GROWTH OF EUCALYPTUS GLOBULUS T. Burgess and N. Malajczuk CSIRO Diviskm ~" F(~restry and Forest Pr~xlucts Perth, Wcsleru Australia ABSTRACT Mycorrhi'zal inoculation offopcn-l~l}inatcd families of Eucalz0ttts Jt.k~bulusreduced the phenotypic variability ol seedlings growth compared with those uninoculated in glasshouse experiments. Seedling growth promotion was in excess d 15t)% for all three ectomyeorrhb'a| fungi tested.
INTRODUCTION Planting of trcc species for timber and pulp in plantations often involves the use of seed collected from the natural range of the species. Scedlots usu',dly represent collections oI a number of half-sibling families and often a number o( pr~wcnances. This genetic variability in scedlots is one of the fact~rs contributing to variable grov,~h ~ff trees in plantation forestry. Lack of the appropriate mycorrhi~,'.al fungi and l~x.aliscd changes in soil conditions are other factors c(mtributing to gr(~vlh disparities in plantations. Ectomycorrhizal fungi are known to stimulate trcc growth and ameliorate extreme variability such as tree death and very p~mr growth of survivors in a wide range ol soil condi6ons, however, the remaining variability in growth has been attributed to the edaphic conditions (Perry et al., 1987; Marx, 1977). To date, the pt)ssibilily that ectomyeorrhizal fungi may reduce growth differences resulting from genetic variability of the planting sh:,ck has not bccn considered. E. globulus is a common plantation species grown in temperate areas of Australia and extensively overseas. Seedlings of E. globulus raised in nurseries represent sccdlots of open-pollinated families from a range ol prt~enances.
The objcclb, c of Ihis sludy was to examine growth and cctomycorrhizal development on
seedlings ~ff 12 open-pollinated families of E.globulus from three provenances. Plants wcrc grown under con~,tz*nt cdaphic condilions in Ihc glasshouse.
4~
MATERIALS AND METHODS
Seedlings of 12 open-pollinated families of E.glohulus representing 3 provenances ((;ceveston. Scamand~r. King Island) wcrc germinated aseptically and inoculated with pure cultures of three fungi; Laccaria laccala Selerodcrma vcrrucosum and Sctchelliogastcr sp. Control seedlings did not receive inoculum After 14 day~, incubatitm with the fungi, seedlings wcrc planted in the glasshouse into pots containing steam pa~.tcuri',td yellow sand supplemented with basal nutrients and ph~sphorus to produce approximately 32)"; maximum grov.lh tor the species. Seedling heights wcrc measured at 2 week intcr~,als and harvested 150 days after planting.
RESULTS AND DISCUSSION
Mycorrhizal dcvelopmcnt attributed to the thrcc fungi was unifi~rmly high (Table 1) and sccdling growlh was promoted abo~.'c thai of unint~culatcd seedlings by an average of 150tTr, for Sclerodcrma vcrrucosum. 171)';" for Sclchclliogastcr sp. and 22tWb for Laccaria laccata.
While variation in biomass was largely dctcrmincd at the opcn-p,.dlinatcd family Icvcl, the rclati',e growth rate, carbohydrate partitioning and phosphorus accumulation appeared to be determined by provenance. At 38 days seedlings of the (;ccvcston provenance wcrc largcst and they had thc greatest root/shoot ratit} while those t~f King Island Wcrc smallest. However at 151}days, King Island prtwenancc had the greatest relative growth rate,
phosphorus uptake and mycorrhizal root length and Gee'.cston the smallest. Scamander
provenance was found to be intermediate between the other two provenances (Table 1).
There was considcrablc variation within and among open-pollinated families in the growth of uninoculatcd seedlings indicating gcnctical variability, This variation in host plants was reduced by inoculatitm with cciomycorrhizal fungi (Figure 1 & 2). The coefficient of variation for uninoeulated plants is 45'::,, while t~r in~culatcd plants this is reduced to 171;. The mode of c~mpcnsatitm is unclear, ht~'.'.cvcr rcsulls ~,upport obser,,ations ~f Marx and Brwm ( 1~71 ) who I'~mnd thai collcctitms tff slash pine varied in their susccptibiliD t~, col~mizalitm and stimulation by fungal s',mbiont~,. At 1511days the seedlings of E. globulus from King Island pr~\cnance had the greater,t m,,corrhi/al r~ot length, indicating they were the mo~,t compalible h~r m~c~r rhi/al dc'.elopment
43
Table I, (;rowl.h chur-',ctcrislics and mycorrhizal development of inoculated and uninoculalcd sccdlings o( Eucalvnlus ulobulus from three provenances, Values in a row with a common subscript do not differ signifi-cantly-(p<0.0$) by a D-ncan's |-test.
PROVENANCE
(;eeveston
Scamander
KinR Island
(;ro,a,lh Churucleristics Inoculated ,~k'edllnMs Total Biomas~, (g) Finc Root/ Coarse Rool Rool/Shtx)t
R(;R (ram/ l{N)mm/Day) % Mycorrhizal Mycorrhizal Root Lcnglh (m) Total R~x)I Lcnglh (m) P Uplakc ( m g / g Rools)
11.00 a
12.50 a
12.71 a
0.32 a 0.44 a
11.32 u I).42 b
0.35 a 0.37 c
2.02 a ¢,1.~1 a
2.10 b ¢)4.32 a
2.28 c 63.65 a
~5.50 a
103.20 b
l(~.b~l b
I.~.ll) a
158.40 b
171.~X1 h
I.U4; a
2.03 u
2.55 b
5.??~ a
4,80 ab
3.63 h
UnlmN.'ululedSeedllnp Total Biomass (g) Fine R(x~l/ Coarse Root R(x)t/Shoot
0.16 a
0.17 a
11.14 a
0.43 a
0..30 b
O..~ b
I(~)mm/Day) Myc(wrhizal
I,~'~ a
I.(~J a
1.74 a
Root Length (m) Tolal Root I~nglh (m) P Uptake
().IX) a
(I.iKJ a
0.(k) a
.~.,~) a
39.YdJ a
31.40 a
1,00 a
1.93 a
1.79 a
R{iR (ram/
( m g / g Roots)
/,4
Setchelliogaster 300
Scmander 3 _, King Island 3 h Geeveston 3
250
Control Geeveston 3
200 E E
Control Scmander 3
/~////::.
150
/./,L
O) <9
//
Control King Island 3
/ / , , . / .."
100
50
0
I,
I
I
i
J
38
66
94
122
150
D~ys after planting
Figure 1. (;rc~vth hi" Eqle,d!vp*,u~ ~10bulu$ from Ihre¢ proven~nceg a|'~er inoculation with Setche I oF'.~ptcr 8p.
compared with ~nlnc,¢ulm[ed plant~;.
45
F i g u r e 2. Growth o f E u ~ l ) ~ t u s g l o b u l u s s e e d ; rwpresentir4[ f o u r openp o l l i n a t ~ f a m i l i e s from t h e King Islm~d p r o v e n m ~ e i n o c u l a t e d ~'ith ~ria ]accata (EIOO4), ~¢lerodezma verrucosUm (HI036) and Setchellio~mter sp, (HIO23) compared with u~inoculated seedlings.
46
CONCLUSION Mycorrhizal im~culation appears tt) reduce the phenotypic variation amongst seedlings~
If this effect is
conlnltln, then nursery inoculation of euealypl seedlings with selected eetomycorrhizas would be of added cconomlcal benefit in plantation forestry. ACKNt)WLEI)GEMENT This research was supported by the W.A. Chip and Pulp Company. Drs 1. Tommerup and T.Grove critically re',ic',~ed the manuscript.
REFERENCES Marx, D.H. and Bryan, W.C. (1';71) Formation of ectomycorrhizae
on
half-sib progenies of slash pine in
aseptic culture. Forest Sci. 17:488-4q2. Marx, D.H., Bryan, W.C. and Cordell, C.E. (1977) Survival and growth of pine seedlings with Pisolithus ectomycorrhizae after two years on reforestation sites in North Carolina and Florida. Forest Sci. 2~,:?,63373.
•
Perry. D.A., Molina, R. and Amaranthus, M.P. (1987) Mycorrhizae, mycorrhb, as, and reforestation: current knowledge and research needs. Can. J. For, Res. 17:929-940.
"Burgess, T. and Ma]ajczuk, N., 198q: The effect of ectomycorrhizal fungi on reducing#~,variatJo,~ of seedling growth of Eucalyptus globulus. Agric., Ecosystems Environ., 28: 41-46.
Agriculture, Ecosystems and Environment, 28 ( 1 9 8 9 ) Elsevier Science Publishers B.V., Amsterdam Printed in Czechoslovakia
41-46
THE EFFECT OF EC'TOMYCORRHIZAL FUNGI ON REDUCING THE VARIATION OF' SEEDLING GROWTH OF EUCALYPTUS GLOBULUS T. Burgess and N. Malajczuk CSIRO Diviskm ~" F(~restry and Forest Pr~xlucts Perth, Wcsleru Australia ABSTRACT Mycorrhi'zal inoculation offopcn-l~l}inatcd families of Eucalz0ttts Jt.k~bulusreduced the phenotypic variability ol seedlings growth compared with those uninoculated in glasshouse experiments. Seedling growth promotion was in excess d 15t)% for all three ectomyeorrhb'a| fungi tested.
INTRODUCTION Planting of trcc species for timber and pulp in plantations often involves the use of seed collected from the natural range of the species. Scedlots usu',dly represent collections oI a number of half-sibling families and often a number o( pr~wcnances. This genetic variability in scedlots is one of the fact~rs contributing to variable grov,~h ~ff trees in plantation forestry. Lack of the appropriate mycorrhi~,'.al fungi and l~x.aliscd changes in soil conditions are other factors c(mtributing to gr(~vlh disparities in plantations. Ectomycorrhizal fungi are known to stimulate trcc growth and ameliorate extreme variability such as tree death and very p~mr growth of survivors in a wide range ol soil condi6ons, however, the remaining variability in growth has been attributed to the edaphic conditions (Perry et al., 1987; Marx, 1977). To date, the pt)ssibilily that ectomyeorrhizal fungi may reduce growth differences resulting from genetic variability of the planting sh:,ck has not bccn considered. E. globulus is a common plantation species grown in temperate areas of Australia and extensively overseas. Seedlings of E. globulus raised in nurseries represent sccdlots of open-pollinated families from a range ol prt~enances.
The objcclb, c of Ihis sludy was to examine growth and cctomycorrhizal development on
seedlings ~ff 12 open-pollinated families of E.globulus from three provenances. Plants wcrc grown under con~,tz*nt cdaphic condilions in Ihc glasshouse.
4~
MATERIALS AND METHODS
Seedlings of 12 open-pollinated families of E.glohulus representing 3 provenances ((;ceveston. Scamand~r. King Island) wcrc germinated aseptically and inoculated with pure cultures of three fungi; Laccaria laccala Selerodcrma vcrrucosum and Sctchelliogastcr sp. Control seedlings did not receive inoculum After 14 day~, incubatitm with the fungi, seedlings wcrc planted in the glasshouse into pots containing steam pa~.tcuri',td yellow sand supplemented with basal nutrients and ph~sphorus to produce approximately 32)"; maximum grov.lh tor the species. Seedling heights wcrc measured at 2 week intcr~,als and harvested 150 days after planting.
RESULTS AND DISCUSSION
Mycorrhizal dcvelopmcnt attributed to the thrcc fungi was unifi~rmly high (Table 1) and sccdling growlh was promoted abo~.'c thai of unint~culatcd seedlings by an average of 150tTr, for Sclerodcrma vcrrucosum. 171)';" for Sclchclliogastcr sp. and 22tWb for Laccaria laccata.
While variation in biomass was largely dctcrmincd at the opcn-p,.dlinatcd family Icvcl, the rclati',e growth rate, carbohydrate partitioning and phosphorus accumulation appeared to be determined by provenance. At 38 days seedlings of the (;ccvcston provenance wcrc largcst and they had thc greatest root/shoot ratit} while those t~f King Island Wcrc smallest. However at 151}days, King Island prtwenancc had the greatest relative growth rate,
phosphorus uptake and mycorrhizal root length and Gee'.cston the smallest. Scamander
provenance was found to be intermediate between the other two provenances (Table 1).
There was considcrablc variation within and among open-pollinated families in the growth of uninoculatcd seedlings indicating gcnctical variability, This variation in host plants was reduced by inoculatitm with cciomycorrhizal fungi (Figure 1 & 2). The coefficient of variation for uninoeulated plants is 45'::,, while t~r in~culatcd plants this is reduced to 171;. The mode of c~mpcnsatitm is unclear, ht~'.'.cvcr rcsulls ~,upport obser,,ations ~f Marx and Brwm ( 1~71 ) who I'~mnd thai collcctitms tff slash pine varied in their susccptibiliD t~, col~mizalitm and stimulation by fungal s',mbiont~,. At 1511days the seedlings of E. globulus from King Island pr~\cnance had the greater,t m,,corrhi/al r~ot length, indicating they were the mo~,t compalible h~r m~c~r rhi/al dc'.elopment
43
Table I, (;rowl.h chur-',ctcrislics and mycorrhizal development of inoculated and uninoculalcd sccdlings o( Eucalvnlus ulobulus from three provenances, Values in a row with a common subscript do not differ signifi-cantly-(p<0.0$) by a D-ncan's |-test.
PROVENANCE
(;eeveston
Scamander
KinR Island
(;ro,a,lh Churucleristics Inoculated ,~k'edllnMs Total Biomas~, (g) Finc Root/ Coarse Rool Rool/Shtx)t
R(;R (ram/ l{N)mm/Day) % Mycorrhizal Mycorrhizal Root Lcnglh (m) Total R~x)I Lcnglh (m) P Uplakc ( m g / g Rools)
11.00 a
12.50 a
12.71 a
0.32 a 0.44 a
11.32 u I).42 b
0.35 a 0.37 c
2.02 a ¢,1.~1 a
2.10 b ¢)4.32 a
2.28 c 63.65 a
~5.50 a
103.20 b
l(~.b~l b
I.~.ll) a
158.40 b
171.~X1 h
I.U4; a
2.03 u
2.55 b
5.??~ a
4,80 ab
3.63 h
UnlmN.'ululedSeedllnp Total Biomass (g) Fine R(x~l/ Coarse Root R(x)t/Shoot
0.16 a
0.17 a
11.14 a
0.43 a
0..30 b
O..~ b
I(~)mm/Day) Myc(wrhizal
I,~'~ a
I.(~J a
1.74 a
Root Length (m) Tolal Root I~nglh (m) P Uptake
().IX) a
(I.iKJ a
0.(k) a
.~.,~) a
39.YdJ a
31.40 a
1,00 a
1.93 a
1.79 a
R{iR (ram/
( m g / g Roots)
/,4
Setchelliogaster 300
Scmander 3 _, King Island 3 h Geeveston 3
250
Control Geeveston 3
200 E E
Control Scmander 3
/~////::.
150
/./,L
O) <9
//
Control King Island 3
/ / , , . / .."
100
50
0
I,
I
I
i
J
38
66
94
122
150
D~ys after planting
Figure 1. (;rc~vth hi" Eqle,d!vp*,u~ ~10bulu$ from Ihre¢ proven~nceg a|'~er inoculation with Setche I oF'.~ptcr 8p.
compared with ~nlnc,¢ulm[ed plant~;.
45
F i g u r e 2. Growth o f E u ~ l ) ~ t u s g l o b u l u s s e e d ; rwpresentir4[ f o u r openp o l l i n a t ~ f a m i l i e s from t h e King Islm~d p r o v e n m ~ e i n o c u l a t e d ~'ith ~ria ]accata (EIOO4), ~¢lerodezma verrucosUm (HI036) and Setchellio~mter sp, (HIO23) compared with u~inoculated seedlings.
46
CONCLUSION Mycorrhizal im~culation appears tt) reduce the phenotypic variation amongst seedlings~
If this effect is
conlnltln, then nursery inoculation of euealypl seedlings with selected eetomycorrhizas would be of added cconomlcal benefit in plantation forestry. ACKNt)WLEI)GEMENT This research was supported by the W.A. Chip and Pulp Company. Drs 1. Tommerup and T.Grove critically re',ic',~ed the manuscript.
REFERENCES Marx, D.H. and Bryan, W.C. (1';71) Formation of ectomycorrhizae
on
half-sib progenies of slash pine in
aseptic culture. Forest Sci. 17:488-4q2. Marx, D.H., Bryan, W.C. and Cordell, C.E. (1977) Survival and growth of pine seedlings with Pisolithus ectomycorrhizae after two years on reforestation sites in North Carolina and Florida. Forest Sci. 2~,:?,63373.
•
Perry. D.A., Molina, R. and Amaranthus, M.P. (1987) Mycorrhizae, mycorrhb, as, and reforestation: current knowledge and research needs. Can. J. For, Res. 17:929-940.
"Burgess, T. and Ma]ajczuk, N., 198q: The effect of ectomycorrhizal fungi on reducing#~,variatJo,~ of seedling growth of Eucalyptus globulus. Agric., Ecosystems Environ., 28: 41-46.