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ŠUMARSKI LIST 13/2005 str. 231     <-- 231 -->        PDF

IZLAGANJE NA ZNANSTVENOM SKUPU - PRESENTATION AT THE INTERNATIONAL SYMPOSIUM Šumarski list SUPLEMENT (2005). 229-237


UDK 630* 181.3


RASPODJELA SITNOG KORIJENJA U ČISTOJ SASTOJIM
OBIČNE SMREKE U STADIJU LETVIKA


FINE ROOT DISTRIBUTION IN A PURE POLE-STAGE
STAND OF NORWAY SPRUCE


Peter JALOVTAR*


SAŽETAK: U radu se analizira raspodjela biomase sitnog korijenja
obične smreke (korijenje s promjerom manjim od 2 mm) u tlu. Istraživane su
četiri sastojine obične smreke u dobi od 21 godine, koje su nastale umjetnom
obnovom na bivšoj poljoprivrednoj površini. Pri osnivanju sastojina, korištena
su dva početna broja sadnica -2.500 i 5.000 komada sadnica obične
smreke po l hektaru s različitim početnim razmacima sadnje. U ovom se radu
procjenjuju razmaci od 4,0 x 1,0 m, 3,0 x 1,3 m, 2,5 x 0,8 m i 2,0 x 1,0 m. Prisutnost
sitnog korijenja u profilu tla istraživano je na dubini od 40 cm. Najviše
vrijednosti biomase sitnog korijenja (7013 kg/ha) ustanovljene su na plohi
s razmakom od 3,0 x 1,3 m, a najniže na plohi s razmakom od 2,5 x 0,8 m
(3945 kg/ha). Odnos između distribucije biomase sitnog korijenja i mase
odumrlog korijenja i dubine tla može se obraditi s linearnom regresijom.
Smanjenje biomase sitnog korijenja uz rastuće dubine tla je različito, najsporije
je na plohi s razmakom od 4,0 x 1,0 m. Za procjenu raspodjele sitnog
korijenja u vodoravnom smjeru, koristili smo gustoću sitnog korijenja izračunatu
na 100 ml tla. Svi razmaci analizirani su na dubini od 2,5 cm, 15 cm
i 35 cm. Odnos između vodoravne udaljenosti i gustoće sitnog korijenja procijenjen
je pomoću funkcije parabole. Na dubini tla od 2,5 cm, funkcija parabole
ima konveksni oblik, s maksimumom u prostoru između redova stabala.
Kako dubina tla raste, oblik funkcije se pretvara u konkavni, s minimumom u
prostoru između redova. Ova je promjena posljedica svojstava korijenskog
sustava obične smreke i ograničenih mogućnosti njezinog rasta pri različitim
razmacima. Odnos između udaljenosti i gustoće sitnog korijenja nije velik,
zbog općenito visoke varijabilnosti korijenske biomase u tlu.


Ključne riječi: obična smreka, sitno korijenje, biomasa, odumrlo sitno
korijenje


UVOD - Introduction


Dominantna funkcija finog i najfinijeg korijenja, tj. (<2,0mm) manje se istražuje nego arhitektura i morfokorijenja
s promjerom pod 2,0 mm je upijanje vode i geneza korijenih sustava pod raznim uvjetima. Njihovo
otopljenih nutrijenata. Stoga se fino korijenje drži indi- intenzivnije istraživanje potaknuto je zbog spomenute
katorom zdravstvenog stanja i snage rasta drveća i sposobnosti njihove reakcije na okolišne promjene,
šumskih sastojina. Funkcija absorbiranja je uvjetovana Razvo j sitno g korijenja i njihov rast prema različianatomijom
sitnog korijenja. Biomasa sitnog korijenja ti m kategorijama promjera ovisni su o vrsti drveća,


starosti, okolišnim uvjetima, bio-sociološkom položa


* Jng. Peter Jaloviar, Katedra za uzgajanje šuma, j u drveta u sastojini te intraspecifičnoj i introspecifi-
Sumarski fakultet, Tehničko sveučilište, "~ -i i t v ti i /ir\ro\ v
Masarykova 24, 960 53 Zvolen, Slovak Republic CI10J konkurenciji (Ko S11 e r et al. (1 968), K O rO t a e V
e-mail:jaloviar@vsld.tuzvo.sk (1997), Wagenknech t (1960), Herte l (1999),




ŠUMARSKI LIST 13/2005 str. 232     <-- 232 -->        PDF

P. Jaloviar: RASPODJELA SITNOG KORIJENJA U ČISTOJ SASTOJIM OBIČNE SMREKE U STADIJU LETV1KA Šumarski list - SUPLEMENT (2005). 229-237
Kodrik (1997). Razvoj i proizvodnja korijenja u komercijalnoj
šumi pod utjecajem je gospodarenja sastojinom
(Jaloviar 1999). Iako većina ovih čimbenika
nisu pod izravnim utjecajem, volumen krošnje i mikroklimatski
uvjeti u sastojini su pod velikim utjecajem
šumsko-uzgojnih zahvata.


Utjecaj početnih razmaka sadnje na razvoj krošnje i
rast obične smreke j e neupitan (Schmidt -Vo g t 1991,
Korpel & Saniga 1995). Pod pretpostavkom daje
ravnoteža nadzemnih i podzemnih dijelova drveta održavana
zdravim pojedinim stablima (Schinozaki et al.


1964), možemo očekivati da razmaci imaju važan utjecaj
na razvoj finog korijenja i njihovu distribuciju u tlu.


Obična smreka (Picea abies L) tipični je predstavnik
vrsta s plosnatim korijenskim sustavom. Dinamika rasta
finog korijenja je autonomna, tj. njihov rast prestaje u
ljeto, čak i u slučaju odgovarajućih uvjeta vlage. Gustoća
sitnog smrekinog korijenja u tlu vrlo je ekstenzivna.
Potencijal prirasta finog korijenja spada u najniže od
naših običnih vrsta drveća. Pojava fiziološki aktivnog
sitnog korijenja koncentrirana je u horizontu tla A.


PODRUČJE PROUČAVANJA I METODE Study area and methods


1UFRO eksperiment sa sadnjom smreke na razmake
osnovan je na lokalitetu Vrch Dobroč 1981-1982, a plohe
s istraživanim razmacima postavljene su godine 1982,
a sada su stare 21 godinu. Plohe su postavljene na elevaciji
od 890 m do 910 m, nagib je između 8 % i 15 % te
ima zapadni aspekt. Površina svake plohe je 0,4 ha.


Tlo na plohama je pjeskovit ilovasti kambisol razvijen
na kristaličnim stjenama. Prosječna godina oborina
je 900mm.


Površina sadašnjeg eksperimenta s razmacima je prvotno
bilo poljoprivredno zemljište, pa stoga homogenost
okoline tla osigurava jedinstvenu mogućnost komparativnih
proučavanja. Mala varijabilnost okoline tla
važna je prednost, zato što ne prikriva razlike u razvoju


korijenog sustava uslijed npr. različitih uzgojnih zahva


ta, razmaka, itd. Detaljni opis područja istraživanja


može se naći uKorpel &Saniga (1994).
Ovaj rad analizira sljedećih 7 razmaka:
Ploha A, razmak 5H0.8 m, gustoća stabala 2.500 ha"1
Ploha B, razmak 4.0H 1.0 m, gustoća stabala 2.500 ha"
Ploha C, razmak 3.0H. 1.3 m, gustoća stabala 2.500 ha"
Ploha D, razmak 2H2 m, gustoća stabala 2.500 ha"´
Ploha E, razmak 3H0.67 m, gustoća stabala 5.000 ha"
Ploha F, razmak 2.5H0.8 m, gustoća stabala 5.000 ha"1
Ploha G, razmak 20H1.0 m, gustoća stabala 5.000 ha"


Tablica 1. Pregled srednjih vrijednosti temeljnih dendrometričkih varijabli na pokusnim plohama


Table 1 The mean values review of the basic dendrometric variables in sample


ploha razmak (m) dbh (cm) visina (m) dužina krošnje (m) širina krošnje (m) volumen 3 krošnje (mVha)
plot spacing (m) dbh (cm) height (m) crown length (m) crown width (m) volume of 3 crowns (m /ha)
F 2.5 H 0.8 m 14.9 10.8 7.2 1.75 28,848
G 2.0H 1.0m 15.3 10.6 7.1 1.8 30,096
C 3.OH 1.3m 15.1 9.5 7.5 2.30 25,954
B 4.0 H 1.0 m 15.5 9.7 7.7 2.45 30,235
D 2.0 H 2.0 m 14.2 10.4 7.0 2.53 28,619
E 3.0 H 0.67 m 14.9 10.2 7.5 2.25 58,742
A 5.0 H 0.8 m 14.5 9.1 7.1 2.30 24,569


Uzorci za procjenu biomase sitnog korijenja uzeti
su prema sustavnom uzorkovanju. U svakom je razmaku
označena linija, na kojoj su u razmacima od 50 cm
uzeti uzorci tla na dubini od 40 cm. Na plohi B8 uzeto
je 8 uzoraka, a na svakoj drugoj plohi po 7 uzoraka.
Uzorcima se rukovalo prema metodologiji spomenutoj,
npr., u djelu Mur ach a (1984).


Uzorci su izvađeni pomoću šupljeg svrdla unutrašnjeg
promjera od 80 mm i duljinom šupljeg dijela od
200 mm.


Stanje (vitalnost) većine finog korijenja može se vidjeti
na mikroskopskim oznakama, a u upitnim slučajevima
rabljen je dvogled povećanja 15 x ili 30 x.


Nakon sušenja korijenja pri 70EC, težina obih kategorija
(vitalne i mrtve) je kvantificirana. Dobivene vrijednosti
su konvertirane u težinu u kg na površini od
1 ha, ili u vrijednosti gustoće sitnog korijenja u mg od
100 ml tla.




ŠUMARSKI LIST 13/2005 str. 236     <-- 236 -->        PDF

PRESENTATION AT THE INTERNATIONAL SYMPOSIUM
Šumarski list - SUPI.EMENT (2005), 229-237


FINE ROOT DISTRIBUTION IN A PURE POLE-STAGE
STAND OF NORWAY SPRUCE


Peter JALOVIAR*


SUMMARY: The paper analyses the distribution of Norway spruce fine
root (roots with diameter under 2 mm) biomass in the soil. Four Norway
spruce stands of the age 21, which were established by artificial regeneration
on former agricultural area have been researched. By the establishment of the
stands two initial numbers of seedlings -2500 and 5000 pes. of Norway
spruce on 1 hectare and various initial spacings have been used. In this paper
the spacings 4,0x 1,0 m, 3,0x 1,3 m, 2,5x 0,8 m and2,0x 1,0 m are evaluated.
The presence of fine roots in the soil profile has been investigated to the
depth of 40 cm. The highest value of fine root biomass (7013 kg/ha) was found
out on the plot with the spacing 3,Ox 1,3 m, the lowest value had the plot with
the spacing 2,5 x 0,8 m (3945 kg/ha). The relation between the distribution of


fine root biomass and necromass and the soil depth could be approached with
a linear regression. The decrease of the fine root biomass with the increasing
soil depth is different, the slowest is on the plot with the spacing 4,0 x 1,0 m.
For the assessment of the fine root distribution in the horizontal direction the
fine root density calculated on 100 ml of soil has been used. All spacings were
analysed in the depth 2,5 cm, 15 cm a 35 cm. The relation between horizontal
distance and fine root density was estimated with the parabola function. In
the soil depth of 2,5 cm the parabola function has a convex shape with a maximum
in the space between the tree rows. With the increasing soil depth the
function shape is turning into a concave one, with the minimum in the space
between the rows. This change is caused by on the root system characteristics
of Norway spruce and on the limited possibilities of his growth by the various
spacings. The relation between the distance and fine root density is not strong
because of generally high root biomass variability in the soil.


Keywords: Norway spruce, fine roots, biomass, necromass


INTRODUCTION


The dominant function of fine and finest foots, i.e. der various conditions. The impulse to more intensive
the roots with the diameter under 2.0 mm is the uptake research of fine roots was their mentioned ability to
of water and dissolved nutrients. Therefore the fine response on the environment changes.


roots are considered a responsive indicator of health The fine root development as well as the growth of
state and growth potential of trees and forest stands. particular diameter categories of roots is impacted esThe
absorbing function of fine roots is conditioned by pecially by tree species, age of tree, environment contheir
anatomy. The fine root biomass production (roots ditions, biosociologicai position of the tree in the


< 2.0 mm diameter) is investigated less than the issue
stand, intraspecific and introspecific competition
of architecture and morphogenesis of root systems un-
Köstle r etal. (1968), Korotaev (1997), Wagcn knecht
(1960), Hertel (1999), Kodrik (1997).


*
Ing. Peter Jaloviar, PhD, Department of Silviculture, The root development and production in commercial
Forestry Faculty, Technical University, Masarykova 24,
forest is influenced besides mentioned factors crucia


960 53 Zvolen, Slovak Republic,


lly by the management of the stand from its origin to


e-mail: jaloviar@vsld.tuzvo.sk




ŠUMARSKI LIST 13/2005 str. 233     <-- 233 -->        PDF

P. Jaloviur: RASPODJELA SITNOG KORIJENJA U ČISTOJ SASTOJINI OBIČNE SMREKE U STADIJU LETVIKA Šumarski list SUPLHMENT (2005). 229-237
REZULTATI I RASPRAVA - Results and discussion


Usporedba određenih razmaka prema biomasi sitnog korijenja


The comparison of particular spacings according to fine root biomass


Proizvodnja biomase sitnog korijenja prema razmacima
bila je prema očekivanju. Uglavnom je nađeno vitalno
sitno korijenje na plohi E s razmacima 3.0 H 1.3 m


(8.817 kg.ha"´).Ova je vrijednost prilično visoka, posebice
u usporedbi s objavljenim podacima za zrele sastojine.
Nakon usporedbe sa sastojinama ostalih vrsta drveća
u istoj fazi rasta, dobiveni podaci se ne mogu držati
ekstremnim. Nađeno je sitno korijenje na plohi F s razmacima
2.5 H 0.8 m (3.945 kg/1).
Težine mrtve mase nisu tako velike kao težine biomase,
pa stoga ne utječu na apsolutne razlike u ukupnim
težinama obih kategorija sitnog korijenja.


U pregledu biomase vitalnog sitnog korijenja (Casper
& Jackson 1997), vrijednosti se kreću od 200 do


5.000 g/m", tj. nakon konverzije s 2000 kg/ha na
50.000 kg/ha. Međutim, autori su uzeli u obzir samo
zrele šumske sastojine. Korotaev (1997), koji je proučavao
biomasu sitnog korijenja u dvije 80-godišnje
sastojine na ilovastim i pjeskovitim tlima, ustanovio
je za pjeskovita tla biomasu od 5.650 kg/ha. Na ilovastim
tlima je ta vrijednost 359 g/m2, što predstavlja


3.590 kg/ha u gornjih 20 cm tla. Kodrik (1998) je
našao 430, odnosno 230 g biomase sitnog korijenja na
1 m~ na dvije plohe s različitim utjecajima imisije u
Moravsko-sliezske Beskydy, što čini 4.300 kg/ha odnosno
2.300 kg/ha. Proučavao je sloj na dubini od
30 cm. Težina mrtve mase bila je 175 g/m" na plohi pod
utjecajem imisija. Prosječna gustoća sitnog korijenja
varirala je s oko 3.000 kg/ha u zrelim sastojinama četinjača
u središnje-europskim uvjetima. Kodrik (1997)
također nalazi vrijednost konstantne biomase (suha
masa) korijenja s promjerom ispod 5 mm od samo
1.300 kg/ha u prašumi rezervata biosfere Pol´ana. S u rovy
(2000) određuje 4.525 kg sitnog korijenja na
1 ha za zaštitnu šumu smreke (zrele sastojine zajedno s
drugim slojem sljedeće generacije).
Tablica 2. Ukupna težina sitnog korijenja prema kategorijama i razmacima


Table 2 Total fine root weight according to the categories and particular spacings


ploha (razmak)


plot (spacing)


A 5.0 H 0.80 m
B 4.0 H 1.0 m
C3.0H 1.3 m
D 2.0 H 2.0 m
E 3.0 H 0.67 m
F 2.5 H 0.8 m
G 2.0 H 1.0 m


biomasa (kg/ha)


biomass (kg/ha)
7572.4
5913.1
7103.0
6744.4
8817.3
3945.2
5066.2


Odnos između dubine tla i biomase vitalnog sitnog
korijenja ispitivanje primjenom metode linearne korelacije.


mrtva masa (kg/ha)


necromass (kg/ha)


1956.6


929.0
1825.1
1190.4
1274.3
1050.4
1146.0
ukupno (kg/ha)


total (kg/ha)


9529.0
6842.1
8928.1
7934.9
10091.7
4995.6
6212.2


Razlike nagiba linija odražavaju različitu distribucije
sitne korjene biomase. Obrazloženje toga mogu biti širi
i asimetričniji razmaci s niskim početnim brojem biljaka


Tablica 3. Temeljni parametri regresije i korelacije odnosa između biomase sitnog korijenja i dubine tla na
svim istraživanim razmacima


Table 3 Basic regression and correlation parameters of the relation between fine root biomass and
soil depth in all researched spacings


ploha (razmak) -plot (spacing)
A(5.0H 0.80 m)
B(4.0H 1.0 m)
C(3.0H 1.3 m)
D (2.0 H 2.0 m)
E (3.0 H 0.67 m)
F (2.5 H 0.8 m)
G(2.0H 1.0 m)


a b r r
2380.9** 57.1* 0.884 0.781
1152.4** 12.0** 0.279 0.077
1419.0** 17 j* * 0.558 0.311
1996.6** 39.7 0.779 0.608
2176.9* 27.2 0.471 0.222
1041.1** 27.9** 0.865 0.748
1190.7** 24.9** 0.578 0.334


(2.500 ha"1), npr. razmak 5,0 x 0,8 m, a razmak sastojine se krošnje drveća ne dotiču u međuprostorima. Stoga
za razvoj krošnje nije smatran tako dobrim kao drugi si-distribucija biomase odgovara manje ili više načinu na
metričniji razmaci. Ovaj je razmak jedini za sada, gdje koji korijenje uzima profil tla u sastojinama s niskom


ŠUMARSKI LIST 13/2005 str. 234     <-- 234 -->        PDF

P. Jaloviar: RASPODJELA SITNOG KORIJENJA U ČISTOJ SASTOJINI OBIČNE SMREKE U STADIJU LETVIKA Šumarski list - SUPI.EMENT (2005). 229-237
gustoćom sastojine, gdje je konkurencija susjednog
drveća manja nego u sastojinama s punom gustoćom. U
simetričnim razmacima raspoloživ prostor sastojine je
potpuno zauzet, a konkurencija sloja krošnje je mnogo


jača nego u prethodnom slučaju. Dokaz za to je vidljivo
intenzivno sušenje u donjim dijelovima krošnje u ovim
razmacima, i stoga se postepena redukcija kapaciteta
krošnje uspoređuje s razmakom 5,0 x 0,8 m.


Usporedba određenih razmaka prema gustoći sitnog korijenja


The comparison of particular spacings according to fine root density


Gustoća sitnog korijenja dana je u mg korijenja u
100 ml finog tla. Prednost ovog parametra je njegova
nezavisnost od debljine istraživanog sloja. U početku
je gustoća sitnog korijenja sa 7 ploha anlizirana kao


cjelina, tj. odnos između prosječne koncentracije i dubine
tla je kvantificirana. Usporedba je izvedena uporabom
metode linearne korelacije.


Vitalno sitno korijenje - Vital fine roots


Pregled koeficijenata regresijskih linija za sve razkoeficijenata
(t-test), ustanovljena je visoko signifimake
danje u Tablici 4. kantna razlika obih ^-koeficijenata s plohe B. Unatoč
signifikantnim razlikama od nule, regresijski koefici


Prema Tablici 4 proizlazi da su svi apsolutini i re


jenti linija s drugih ploha ne razlikuju se signifikantno


gresijski koeficijenti, koji su determinantni za oblik


jedni od drugih.


određene relacije, statistički visoko signifikantni različito
od nule. Ispitivanjem razlika između regresijskih


Tablica 4. Temeljni regresijski i korelacijski parametri odnosa između prosječne gustoće sitnog korijenja i
dubine tla na svim istraživanim razmacima


Table 4 Basic regression and correlation parameters of the relation between average fine root density
and soil death in all researched spacings


ploha (razmak) -plot (spacing)
A (5.0 H 0.80 m)
B(4.0H 1.0 m)
C(3.0H 1.3 m)
D (2.0 H 2.0 m)
E (3.0 H 0.67 m)
F (2.5 H 0.8 m)
G(2.0H 1.0 m)


a b s xb r 2
r
370.2** 10.10* 3.41 0.83* 0.68
267.5** 6.09** 1.21 0.61** 0.37
332.6** 8.96** 1.39 0.73** 0.53
256.8* 5.41 3.94 0.55 0.32
154.3* 3.52 1.98 0.66 0.44
257.5** 8.55** 0.98 0.81** 0.65
286.5** 8.71** 1.49 0.68** 0.46


*
B signifikantna razlika od nule, **B visoko signifikantna razlika od nule, parameteri Sbx
i r nisu ispitani na razliku od nule
*
B significant difference from zero, ** B highly significant difference from zero, parameters sbx
and r are not tested to the difference from zero
Korelacije između dubine tla i koncentracije sitA
(5,0 H 0,8) i F (2,5 H =,8 m), dok izračunata linija
nog korijenja prilično su slabe, najjača je na plohama iznosi 69 % odnosno 62 % varijabilnosti.


Odumrlo sitno korijenje Dead fine roots


I u ovom je slučaju primijenjena metoda linearne
korelacije za usporedbu odnosa između koncentracije
sitnog korijenja i dubine tla. Rezultati su prikazani u
Tablici 5.


Jačina korelacija je varijabilnija na odumrlom korijenju
nego vitalnom. Iz prijašnjeg se iskustva zna kada
odumrlo sitno korijenje pokazuje bitno veću varijabilnost
i često nikakav odnos s dubinom tla.


Na plohama s većim početnim brojem biljaka na
1 ha, pad koncentracije biomase kao i ukupne sitne rizomase
s povećanjem dubine tla manji je nego na plohama
s manje gustim razmacima. Očekivaniji rezultat je, da će


koncentracija finog korijenja biti veća pri većoj gustoći
i simetričnijim razmacima, što će se potvrditi manje izraženim
gradijentima dubine. Možemo pretpostaviti da
je u usporedbi s ljetom vlaga tla bila viša već na početku
jesenskog rasta sitnog korijenja, u vrijeme njihove
sječe. Na plohama s gušćim razmacima je intercepcija
svakako veća, a slojevi tla ispod 20 cm su očigledno
suhlji u vrijeme sječe nego na plohama s manje gustim
razmacima, s manjim stupnjem sklopa. Stoga možemo
pretpostaviti daje na plohama s gustim razmacima rast
korijenja bio ograničen samo na horizontu tla A u vrijeme
sječe, tj. na slojevima na dubini 0-20 cm.




ŠUMARSKI LIST 13/2005 str. 235     <-- 235 -->        PDF

P. Jaloviar: RASPODJELA SITNOG KORIJENJA U ČISTOJ SASTOJIN1 OBIČNE SMREKE U STADIJU LETVIKA Šumarski list - SUPLEMENT (2005). 229-237
Tablica 5. Temeljni regresijski i korelacijski parametri odnosa između prosječne gustoće sitnog korijenja i
dubine tla na svim istraživanim razmacima


Table 5 Basic regression and correlation parameters of the relation between average vital fine root density
and soil death in all researched spacings


Ploha (razmak) - plot (spacing)
A (5.0 H 0.80 m)
B(4.0H 1.0 m)
C(3.0H 1.3 m)
D (2.0 H 2.0 m)
E (3.0 H 0.67 m)
F (2.5 H 0.8 m)
G(2.0H 1.0 m)


a b s xb r r
89.3** 2.32 0.98 0.76 0.58
27.16** 0.25NS 0.19 0.20NS 0.04
51.9* 0.50NS 0.43 0.18NS 0.03
51.9** 1.14* 0.33 0.86* 0.75
93.7** 2.88* 0.94 0.84* 0.70
43 9** 0.99** 0.23 0.55** 0.30
47.0** 0.94NS 0.36 0.38* 0.14


** B visoko signifikantna razlika od nule, NSB bez signifikantne razlike od nule, parameteri Sbx i r2 nisu
ispitani na razliku od nule.
** B highly significant difference from zero, mB no significant dirterence from zero, parameters Shx and r´


are not tested to the difference from zero


ZAKLJUČCI
Različita struktura sastojine uzrokovana različitim
razmacima odražava se na svim istraživanim parametrima.
Ustanovljena je različita proizvodnja biomase
sitnog korijenja iz ukupne proizvodnje rizomase, no to
otkriće se ograničava na razdoblje života sastojine, kada
je čak i u manje gustim razmacima konkurencija
među drvećem u razmacima između redova tako velika,
da dolazi do intenzivnijeg sušenja donjih dijelova
krošanja. Također je potvrđena pretpostavka o različitim
načinima distribucije biomase sitnog korijenja.
Odnos između gustoće sitnog korijenja i dubine tla
predstavlja rzaličite oblike prema razmacima.


LITERATURACasper, B. B.,R. B.Jackson, 1997: Plant competition
underground. Annu. Rev. Ecol. Syst. 28:
545-570
Hertel , D., 1999: Das Feinwurzelsystem von Reinund
Mischbeständen der Rotbuche: Struktur,
Dynamik und interspezifische Konkurrenz. Dissertationes
Botanicae. Bd317, 187 s.
Jaloviar, P, 1999: Produkcia jemnych korenov v
röznych typoch smrekovych porastov. In: Atmosfera
21. Storocia, organizmy a ekosystemy.
TU Zvolcn 94-97.
Kodrik , M., 1997: Vyskum podzemnej biomasy
smreciny (korenov) a jej dlzky v BR Pol´ana. In:
Biosfericke rezerväcie na Slovensku, 133-137.
Kodrik , M., 1998: Investigation of fine roots of Picea
abies ecosystem in the northern Slovakia.
Ekologia 17 (4): 358-363.
Korotacv,A . A, 1997: Wurzelmorphologische Untersuchungen
der Fichte (Picea abies (L.)
Karst.) auf Sand- und Schluffboden im Gebiet
von St. Peterburg. Forstarchiv 68 (3): 102-108.


Conclusions
Kako bi se proširilo znanje rizologije šumskog drveća,
bit će potrebno postaviti fokus na slična proučavanja,
posebice na istraživanje problema promjena u
proizvodnji sitnog korijenja u šumskim sastojinama.
Doprinos ovog rada je također u tome da može biti dobar
temelj za planiranje sljedećeg istraživanja sustava
korijenja u pokusu s razmacima u Vrch Dobroč, jer
pruža pouzdane podatke o varijabilnosti sitnog korijenja
i osnovne uzorke njihove distribucije u prostoru tla.


- References
Korpel´, Š., M. Saniga, 1995: Vplyv rozdiclneho
počtu sadenic a ich sponu na rast a formovanie
smrekovych porastov. Vedecki Studie 3. TU Zvolen,
39 s.


Köstler, J. N., E. Brückner, H. Bibelriether,
1968: Die Wurzel der Waldbäume. Paul-Parey-
Verlag. Berlin, Hamburg 282 s.


M u r a c h, D., 1984: Die Reaktion der Feinwurzel von
Fichte (Picea abies Karst. L.) auf zunehmende
Bodenversauerung. Göttinger Bodenkdl. Ber.
77:1-126.


Schmidt-Vogt, EL, 1991: Die Fichte. Bd. H/1. Paul-
Parrey-Verlag. Hamburg, München, 105-106.


Surovy, P, 2000: Porovnanie produkeie biomasy
jemnych korenov v lesnych porastoch s prevahou
smreka obhospodarovanych podrastovym a
vyberkovym hospodärskym sposobom. Diplotnovä
praca, LF Tu vo Zvolene. 36 s.


Wagenknecht ,
E., 1960: Beiträge zur Kenntnis der
Wurzelausbildung verschiedener Bestockungen.


Mitteilungen der Staatforstverwaltung Bayerns.




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P. Jaloviar: FINE ROOT DISTRIBUTION IN A PURE NORWAY SPRUCE POLE-STAGE STAND Šumarski list SUPLEMENT (2005), 229-237
the emergence of next forest generation (Jaloviar Norway spruce {Picea abies L. Karst.) is a typical
1999). The most of these factors can´t be affected direpresentative
of the tree species with the flat root sysrectly,
but e.g. the crown volume and microclimatic tem. The growth dynamics of its fine roots is autonoconditions
in the stand are decisive impacted by silvi-mous, i.e. their growth stops in the summertime also in
cultural interventions as well. the case of appropriate moisture conditions. The densi


The influence of the initial spacing on the crown dety
of spruce fine roots in the soil is very extensive. The
velopment and the growth of Norway spruce is indispuincrement
potential of its fine roots belongs to the lotable
(Schmidt-Vogt 1991, Korpel ´ & Saniga west between our common tree species. The appearan1995).
Assuming that the balance between the above-ce of physiological active fine roots is concentrated in


ground and underground part of the tree is kept by the the soil horizon A.
healthy individuals (Schinozaki etal. 1964), we can
expect the spacing has an important influence on the fine
root development and distribution in the soil as well.


STUDY AREA AND METHODS


The IUFRO spruce plant spacing experiment was plot D, spacing 2H2 m, stem density 2,500 ha"´
established on locality Vrch Dobroč in 1981-1982, the


plot E, spacing 3H0.67 m, stem density 5,000 ha"1


plots with investigated spacings were established 1982


plot F, spacing 2.5H0.8 m, stem density 5,000 ha"1


and at present they are 21 years old. The plots are located
in the elevation from 890 to 910 m, the slope ranges plot G, spacing 2.0H1.0 m, stem density 5,000 ha"´
from 8 to 15 % and it has west aspect. The area of each The samples for estimation of fine root biomass were
plot is 0.4 ha. taken according to systematic sampling. In each spacing


The soil on the plots is sandy loam cambisol that a line was marked, where in the distance of 50 cm the
developed on crystalline rock. Average annual precipisoil
cores up to the depth 40 cm were taken. On the plot
tation is 900 mm. B 8 probes were established, on each other plot 7 pro


bes. The samples were handled according to the metho


The area of present spacing experiment was agri


dology mentioned e.g. in the work of M u r a c h (1984).


cultural land initially and therefore the homogeneity of
the soil environment provides an unique possibility for The samples (soil cores) were taken by a hollow aucomparative
studies. The low variability of the soil enger
with the inner diameter 80 mm and the length of the
vironment is an important advantage, because it don´t hollow part 200 mm.
conceal the differencies of the root system developThe
state (vitality) of the most fine roots can be
ment due to e.g. various silvicultural interventions, identified from the macroscopic marks, in the questiospacing
etc. Detailed description of the research area nable cases a binocular glass with the magnification
can be found inKorpel´&Saniga (1994). 15x or 30x was used.


This paper analyses following 7 spacings: After the drying of roots at 70EC the weight of both
categories (vital and dead) was quantified. The acqui


plot A, spacing 5H0.8 m, stem density 2,500 ha"´ plot


red values were converted into the weight in kg on the


B, spacing 4.0H1.0 m, stem density 2,500 ha"1


area of 1 ha or into the values of fine root density in mg
plot C, spacing 3.OH 1.3 m, stem density 2,500 ha"´ of 100 ml of soil.


RESULTS AND DISCUSSION


The comparison of particular spacings according to fine root biomass


The production of the fine root biomass varies acThe
nccromass weights are not so high as the biocording
to the spacing as expected. Mostly vital fine mass weights and therefore they don´t impact the absoroots
were found on the plot E with the spacing lute differencies in the total weights of both fine root
3.OH 1.3 m (8,817 kg.ha"´). This value is quite high, escategories
either.
pecially in the comparison with the data published for In the review of vital fine root biomass compiled by
the mature stands. After the comparison with the stands Casper & Jackson (1997) the values range from 200


of other tree species in the same growth phase, acquito
5,000 g/m , i.e. after conversion from 2,000 kg/ha to
red data can´t be consider extreme. At least vital fine 50,000 kg/ha. However, the authors have considered onroots
were found on the plot F with the spacing 2.5 H ly mature forest stands. Korotaev (1997), who inves


0.8 m (3,945 kg.ha"´). tigated the fine root biomass in two 80 years old stands


ŠUMARSKI LIST 13/2005 str. 238     <-- 238 -->        PDF

P. Jaloviar: FINE ROOT DISTRIBUTION IN A PURE NORWAY SPRUCE POLE-STAGE STAND Šumarski list - SUPLHMENT (2005), 229-237
on loam and sandy soils, states for sandy soils the biomass
of 5,650 kg/ha. For the loam soils he states the
value 359 g/m2, what presents 3,590 kg/ha in the top 20
cm of soil. Kodri k (1998) found out 430 respectively
230 g of fine root biomass on 1 m on two plots with different
immission impact in the region of Moravskosliezskc
Beskydy, what constitutes 4,300 respectively
2,300 kg/ha, he investigated the layer with depth of
30 cm. The necromass weight was 175 g/m2 on the plot
under immission impact. The average fine root density
varies about 3,000 kg/ha in mature coniferous stands in
central European conditions. Kodri k (1997) also states
the value of constant biomass (dry mass) of roots
with diameter under 5 mm only 1,300 kg/ha for the primeval
forest in Biosphere Reserve Pol´ana. Surov y
(2000) gives 4,525 kg of fine roots on 1 ha for a spruce
shelterwood forest (mature stand together with second
layer of next generation).


The relation between the soil depth and vital fine
root biomass was tested using the method of linear correlation.


The comparison of particular spa


The fine root density is given in mg of fine roots in
100 ml of fine soil. The advantage of this parameter is
its independence from the thickness of investigated
layer. At first the fine root density from 7 plots was


Vital f


The overview about the coefficients of regression
lines for all spacings is given in Table 4.


From Table 4 results that all absolute and regression
coefficients, which are determinant for the shape of
particular relation, are statistically high significant different
from zero. By testing of diffcrencies between the
regression coefficients (t-test) a high significant difference
of the b-coefficient from the plot B was found.


The differencics in the incline of the lines reflect the
various distribution of the fine root biomass. The explanation
for this can be that by wider and more asymmetric
spacings with low initial plant number (2,500 ha"´)


e.g. by the spacing 5.0 x 0.8 m, the available stand space
for the crown development is not used as good as by
the other more symmetric spacings. This spacing is the
only one at present, where the crowns of trees don´t
touch in the inter-row space. Therefore the biomass
distribution corresponds more or less to the way how
the roots take the soil profile in the stands with low
stand density, where the competition of neighbour trees
is less than in the stands with full density. In symmetric
spacings the available stand space is fully used and the
competition in the crown layer is much stronger than in
the previous case. The proof of it is the visible more intensive
drying in the lower parts of the crowns in these
spacings and therefore the gradual reduction of the
crown capacity compared to the spacing 5.0 x 0.8 m.
igs according to fine root density


analysed as a whole i.e. the relation between the avera


ge concentration and the soil depth was quantified. The


comparison was conducted using the method of linear


correlation.


i roots


Despite the significant diffcrencies from zero the regression
coefficients of the lines from other plots differ
not significant from each other.


The correlations between the soil depth and fine
root concentration is quite weak, the strongest is on the
plot A (5.0 H 0.8) and F (2.5 H 0.8 m), while the computed
line explains 69 % respectively 62 % of the variability.


Dead fine roots


Also in this case the method of linear correlation
was used for the comparison of relation between the fine
root concentration and the soil depth. The results are
shown in the Table 5.


The strength of the correlations is more variable at
dead roots than vital roots. From previous experiences
the state is known, when the dead fine roots shows substantially
higher variability and often nearly no relation
to the soil depth.


On the plots with higher initial plant number on 1 ha
the decrease of biomass concentration as well as total fine
rhizomass with the soil depth is less strong than on
the plots with less dense spacings. The more expected


result will be that at higher density and symmetric spacing
the fine root concentration will be more homogenous
what will be confirmed by less marked depth gradients.
We can assume that compared with the summertime
the soil moisture was already higher at the beginning
of autumn growth of fine roots optionally in the time
of their harvesting. On the plots with more dense
spacings surely the interception is higher and soil layers
under 20 cm were obvious more dry in the time of the
harvesting than on the plots with less dense spacings
with lower canopy degree. Therefore we can presume
on the plots with dense spacing also the root growth was
limited only on the soil horizon A in the time of harvesting
i.e. on the layers in the depth 0-20 cm.




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P. Jaloviar: FINE ROOT DISTRIBUTION IN A PbRE NORWAY SPRUCH POEH-STAGE STAND Šumarski list - SUPLEMENT (2005), 229-237
CONCLUSIONS


The different stand structure caused by a different
spacing reflects on all investigated parameters. The different
fine root biomass production from the total rhizomass
production was found out, while this finding have
to be limited on the period of stand´s life, when even in
the less dense spacings the competition between the
trees in the inter-row space become to be so high, that it
comes to the more intensive drying of the lower parts of
crowns. The assumption about different way of fine root
biomass distribution was confirmed as well. The relation
between fine root density and soil depth presents
different shapes according to the spacing.


To be able to enlarge the knowledge in the forest
trees rhizology it will be necessary to focus similar studies
especially on the investigation of the issue of changes
in the fine root production of forest stands. The contribution
of this paper is also that it can be a good base
for the planning of next root systems research in the spacing
experiment Vrch Dobroc because it provides a reliable
informations about the fine root variability and basic
patterns of their distribution in the soil space.