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NEEDLE STRUCTURES AND EPIPHYTIC MICROFLORA OF SCOTSPINE (
PINUSSYLVESTRIS
L.) UNDER HEAVY AMMONIADEPOSITION FROM FUR FARMING
JAANA B¨ACK
1
;
2
;?
, MINNA TURUNEN
3
, ARI FERM
4
;??
and SATU HUTTUNEN
1
1
Department of Biology, Botany, Univ. of Oulu, P.O. Box 333, FIN–90571 Oulu, Finland;
2
FinnishForest Research Institute, P.O. Box 18, FIN–01301 Vantaa, Finland;
3
Arctic Centre, Univ. of Lapland, P.O. Box 122, FIN–96101 Rovaniemi, Finland;
4
Finnish Forest Research Institute,Kannus Research Station, P.O. Box 44, FIN–69101 Kannus, Finland
(Received 9 November 1995; accepted 20 July 1996)
Abstract.
Scots pine (
Pinus sylvestris
) needles from forest sites differing in distance from big furfarms emitting large amounts of ammonia and ammonium (=NH
y
) were examined by scanningand transmission electron microscopy. Specific features indicating an ammonium-nitrogen overload,such as an abundance of needle surface organisms and modifications in cellular and wax structures,were classified. Throughout the study area (up to 1000 m from the farms), mesophyll cells hadthin cytoplasm and folding plasmalemma indicating frost damage. Phloem damage attributable toa possible nutrient imbalance was also observed. Chloroplast membranes were undulating and theoccurrence of leaf surface organisms (e.g. aerophilic algae) was more abundant at the closest sites.The changes were related both to the direct effects of dry NH
y
deposition on the needles, and to theeffects operating via soil acidification. The needle epicuticular waxes proved to be structurally ratherinert against the influence of ammonium compounds, since no significant changes due to NH
y
wereobserved.
Key words:
pine needles, ammonium deposition, microscopy, microflora
1. Introduction
Nitrogen is considered a minimum factor in boreal ecosystems, limiting forestgrowth, as is indicated by the increases in biomass growth that frequently followfertilization with nitrogen compounds (Van Dijk and Roelofs, 1988; Kachi andRorison, 1989).
Pinus sylvestris
(L.) is well adapted to acidic soils, and thereforehasastrongpreferencefor ammonium(NH
+
4
) overnitrate (NO
3
) as asoilnitrogensource (Marschner, 1986), but pine forests exposed to excess nitrogen originatingeither from fertilization or atmospheric deposition suffer from many metabolicdisturbancesand from nutrient imbalances in the foliage (Ferm
et al
., 1990; Dueck
et al
., 1991; Pearson and Stewart, 1993; Fangmeier
et al
., 1994). These alterationsmay lead to poor survival during periods of frost or drought and a characteristicexternal appearance following loss of apical dominance and repeated dieback andyellowing of the leader shoots. In addition, the occurrence of a microflora (mainly
?
Correspondence to: Jaana B¨ack, Finnish Forest Research Institute, P.O. Box 18, FIN–01301Vantaa, Finland.
??
Deceased.
Water, Air, and Soil Pollution
100:
119–132, 1997.c
1997
Kluwer Academic Publishers. Printed in the Netherlands.
120
J. B¨ACK ET AL.
algae) on leaf surfaces seems to show a positive correlation with the amount of nitrogen deposition (G¨oransson, 1988; Thomsen, 1992; Van Dijk
et al
., 1992).Depositions of ammonia (NH
3
) and ammonium originate mainly from agricul-turalsources,andthereforearemorepronouncedinruralareas,whilethedepositionof oxidized nitrogen compounds prevails in urban areas. NH
3
in the atmosphere iseither rapidly converted to NH
+
4
in a reaction increasing the pH of precipitation,or subjected to dry deposition. High concentrations of NH
3
in deposition are thusonly found in areas close to emission sources, whereas the proportion of NH
+
4
increases with distance from the source (Fangmeier
et al
., 1994). Deposited NH
+
4
contributes greatly to the acidification of forest soils, increasing the leaching of mineral elements and also nitrate after nitrification, and it also reduces the growthof mycorrhiza (Van der Eerden
et al
., 1992). A shift in forest plant species compo-sition towards more nitrophilous species is commonly found in areas with a highNH
y
(NH
+
4
+ NH
3
) load (see references in Fangmeier
et al
., 1994).Deposited NH
+
4
and NH
3
are absorbed by the canopy principally through thestomata (e.g. Dueck
et al
., 1991), althoughsome absorptionmay also occur via thecuticle after dissolution in the thin water film on needle surface especially in lowpH (Van Hove
et al
., 1989). After entering the intercellular space,both compoundsdissolve in the water films surrounding the mesophyll cells and occur solely asNH
+
4
(Van Hove
et al
., 1987; Fangmeier
et al
., 1994). NH
+
4
can be assimilatedby both the roots and the needles. In chloroplasts it is first converted through theglutamine/glutamate cycle to form either storage compounds or structural proteins(Mitchell and Stocking,1975). Excess availability of NH
+
4
changes the amino acidpattern in conifer needles and leads to a lack of photoassimilates due to increasedenergy demands in ammonium assimilation, especially at low light intensities(Pietil¨a
et al
., 1991; Van der Eerden and Perez-Soba, 1992; Pearson and Stewart,1993).The effects of NH
y
deposition on coniferous forests have mainly been studiedin Central Europe, and especially in the Netherlands, where the emissions arelargest (e.g. Van der Eerden, 1982; Roelofs
et al
., 1985). They have been lesscomprehensively examined under boreal climatic conditions. Our objectives wereto examine whether the earlier observed biochemical alterations in Scots pineneedlesinanareawith intensivefur farming (Ferm
etal
., 1990,Pietil¨a
et al
., 1991)could be attributed to detectable changes in needle surface and cellular structuresin light or electron microscope. These changes may be induced either by a directeffect of exposure to atmospheric NH
y
deposition, or by mineral deficienciesinduced indirectly by long-term changes in soil chemistry.
PINE NEEDLE STRUCTURES AND AMMONIA DEPOSITION
121
2. Material and Methods
2.1. S
ITE DESCRIPTION
The effects of excess NH
y
deposition on the needle structures of Scots pine (
Pinussylvestris
) were examined in seven plots situated in an area with several large furfarms in Kannus, Finnish Ostrobothnia (64
N) (Ferm
et al
., 1990; Pietil¨a
et al
.,1991). The area, regarded as one of the principal fur farming regions in Finlandfrom the early 1970’s onwards, was characterized by a density of 7–10 mink andfox farms km
2
. The field layer of all the forest plots was originally dominated by
Vaccinium myrtillus
L., but after 20 years of continuous high nitrogen depositionit had been substituted by grasses such as
Deschampsia flexuosa
(L.) Trin. on themost exposed plots. The dominant trees were 20–40-year-old planted Scots pines(diameter at breast height 10–20cm). Sevenplots, each including one or two trees,were established at the following distances from the large mink and fox farms:
<
100 m (three plots, five trees), 250–500 m (two plots, three trees), and
>
1000 m(two control plots, four trees).Earlierinvestigationshadrevealedextremelyhighfoliarnitrogenconcentrations(max25mgg
1
dryweight),deficienciesofB(min2.9
gg
1
dryweight)andMg(min0.48mgg
1
dryweight)andaccumulationofproteinsandespeciallyarginineintheneedlesofthesamepinestandsclosetothefur farms(Ferm
etal
.,1990).Thevisible crown damage (bushy growth, bud and needle loss, colour changes in thefoliage) was closely correlated with distance from the fur farms, and an abundantcoverage of green algae on the trunks of the trees was seen in the vicinity of thefarms. Thetotal depositionof NH
+
4
under thecanopyin summer1988rangedfrom0.128 kmol ha
1
in the control areas to 1.805 kmol ha
1
in the closest plots, andfrom 0.050 kmol ha
1
to 0.367 kmol ha
1
, respectively, in an open field. NO
3
deposition in the throughfall was 0.032–0.048 kmol ha
1
near the large farms andless than 0.03 kmol ha
1
in the control areas (Ferm
et al
., 1990). The averageannual temperatures in 1989 and 1990 were 4.7
C and 4.0
C, respectively, thelong-term mean being 3.1
C. The lowest monthly mean temperature in 1990 was–12.8
C (January) and the highest 19.7
C (July-August) (Finnish MeteorologicalInstitute, 1989; 1990). The minimum temperature in 1990 had occurred five daysbefore the sampling (–17.5
C). Annual precipitation in both years was 455 mm,which was 112% of the long-term average.2.2. S
AMPLING AND MEASUREMENTS
Two to three branches per tree were collected with pole-clippers from the middlepart of the canopy of dominant pine trees on November 19th, 1990. Needle yearclass numbers were counted, after which three to five visibly healthy current(c) and previous year (c+1) needles per tree were prepared for scanning electronmicroscopy(SEM)accordingtoTurunenandHuttunen(1991)andfortransmissionelectronmicroscopy(TEM)accordingtoB¨ackandHuttunen(1992).Theremaining
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