The role of stem recharge in reducing the winter desiccation of Picea engelmannii (Pinaceae) needles at alpine timberline

The winter desiccation of needles is thought to limit tree growth and survival within alpine timberline ecotones of the southern Rocky Mountains, USA. To better understand the factors contributing to this desiccation damage, the extent to which stem water was available to needles of Picea engelmannii undergoing desiccation at timberline near Monarch Pass, Colorado, was monitored throughout the winter. Severed shoots experienced significantly greater desiccation than did intact shoots, indicating the availability of stem water to needles despite presumably frozen soil, roots, and stems. A model of water relations during winter predicted more extreme desiccation of severed shoots than observed. This suggests that one or more of the common assumptions concerning the winter water relations of timberline trees is in error. The influence of cold, dry conditions on the cuticular conductance of Picea engelmannii needles is not known and therefore not accounted for in current models of winter water relations. The assumption that cuticular conductance is not influenced by temperature or humidity is a likely source of error in such models.     

Are flowers physiological sinks or faucets? Costs and correlates of water use by flowers of Polemonium viscosum

Water loss through inflorescences may place extreme demands on plant water status in arid environments. Here we examine how corolla size, a trait known to influence pollination success, affects the water cost of flowering in the alpine skypilot, Polemonium viscosum. In a potometry experiment, water uptake rates of inflorescences were monitored during bud expansion and anthesis. Corolla volume of fully expanded flowers predicted water uptake during bud expansion (R ²=0.61, P=0.0375) and corolla surface area predicted water uptake during anthesis (R ²=0.59, P=0.044). To probe mechanisms underlying the relationship between corolla size and water uptake, cell dimensions and densities were measured in several regions of fully expanded corollas. Corolla length was positively correlated with cell length in the middle of the corolla tube and cell diameter in the corolla lobe (Pearson's r from 0.26–0.33, n=86, P ≤ 0.05). Cell density was negatively correlated with cell dimensions in the upper corolla tube and lobe (Pearson's r from –0.39 to –0.42, P ≤ 0.0015). These findings suggest that more water may be required to maintain turgor in large corollas in part because their tissues have lower cell wall densities. The carbon cost of water use by flowers was assessed in krummholz and tundra habitats for P. viscosum flowering, respectively, during dry and wet portions of the growing season. For plants in full flower, average leaf water potentials were significantly more negative (P=0.0079) at mid-day in the krummholz (June) than in the tundra (July), but were similar before dawn (P=0.631). Photosynthetic rate at the time of flowering declined significantly with increasing corolla size in the krummholz (P=0.0376), but was unrelated to corolla size on the tundra (P>0.72). Plants losing water through large corollas may close leaf stomata to maintain turgor. If photosynthesis limits growth in this perennial species, then the water cost of producing large flowers should exacerbate the cost of reproduction under dry conditions. Such factors could select for flowers with smaller corollas in the krummholz, countering pollinator-mediated selection and helping maintain genetic variation in corolla size components of P. viscosum. Received: 5 May 1998 / Accepted: 2 October 1998     

Temporal dynamic of wood formation in Pinus cembra along the alpine treeline ecotone and the effect of climate variables

We determined the temporal dynamic of cambial activity and xylem development of stone pine (Pinus cembra L.) throughout the treeline ecotone. Repeated micro-sampling of the developing tree ring was carried out during the growing seasons 2006 and 2007 at the timberline (1,950 m a.s.l.), treeline (2,110 m a.s.l.) and within the krummholz belt (2,180 m a.s.l.) and the influence of climate variables on intra-annual wood formation was determined. At the beginning of both growing seasons, highest numbers of cambial and enlarging cells were observed at the treeline. Soil temperatures at time of initiation of cambial activity were c. 1.5°C higher at treeline (open canopy) compared to timberline (closed canopy), suggesting that a threshold root-zone temperature is involved in triggering onset of above ground stem growth. The rate of xylem cell production determined in two weekly intervals during June through August 2006–2007 was significantly correlated with air temperature (temperature sums expressed as degree-days and mean daily maximum temperature) at the timberline only. Lack of significant relationships between tracheid production and temperature variables at the treeline and within the krummholz belt support past dendroclimatological studies that more extreme environmental conditions (e.g., wind exposure, frost desiccation, late frost) increasingly control tree growth above timberline. Results of this study revealed that spatial and temporal (i.e., year-to-year) variability in timing and dynamic of wood formation of P. cembra is strongly influenced by local site factors within the treeline ecotone and the dynamics of seasonal temperature variation, respectively.     

Foliar carbon isotope discrimination in Larix species and sympatric evergreen conifers: a global comparison

Larches (Larix spp.), deciduous conifers, occur in the northern hemisphere in cold-temperate and boreal climates – an environment normally thought to favor evergreen tree species. We compare foliar carbon isotope discrimination (Δ), instantaneous water use efficiency, total foliar nitrogen concentration, and specific leaf area (for a subset of sites) between Larix spp. and co-occurring evergreen conifers at 20 sites throughout the natural range of larches. Except for Larix occidentalis in the xeric Intermountain West, USA, Δ is significantly (P < 0.05) greater for larches than co-occurring evergreen conifers at 77% of the sites, suggesting that larches use water less efficiently. At elevations greater than 3000 m, the Δ of Larix spp. and co-occurring conifers converge, suggesting that water is not the limiting resource. Foliar nitrogen concentration and specific leaf area are two ecophysiological characteristics that are positively correlated with high photosynthetic capacity. Foliar nitrogen concentration is significantly greater for larches than evergreen conifers at 88% of the sites and specific leaf area is approximately three times greater for larches than co-occurring conifers. Future studies should examine the potential effect that global warming may have on the distribution of larch forests because the water use efficiency of larches is commonly less than co-occurring evergreen conifers and the boreal and high-latitude environments are likely to experience the greatest climate warming. Received: 23 May 1997 / Accepted: 28 October 1997     

Influence of krummholz mat microclimate on needle physiology and survival

Summary Microclimate and photosynthesis of krummholz mat growth forms of Picea engelmanii (Parry) and Abies lasiocarpa [Hook.] Nutt. were investigated to determine structural features which may aid survival in alpine environments. The structure of krummholz mats was described in terms of the vertical distribution of leaf area index and leaf area density, which exceeded 50 m^-1 (based on total leaf surface area) near the canopy surface and approached zero below 30 cm from the surface in both species. Photosynthetic photon flux density (PPFD, 0.4–0.7 m wavelengths) and wind decreased by an average of 6 and 50-fold, respectively, between 1 m above and 10 cm below mat surfaces in both species. Needle temperatures on a P. engelmannii krummholz mat during July averaged about 2°C above air temperature during the day, with a maximum overtemperature of greater than 20°C above T _air during one sunlit period. At night, needle temperatures averaged 3–4°C below T _air.Net photosynthesis in year-old P. engelmannii shoots reached a maximum at 15–20°C during July and August. Surface shoots were light saturated at near 1200 moles m^-2s^-1 PPFD, and had higher photosynthetic rates than subsurface, predominantly shaded shoots above 800 moles m^-2s^-1. Shade shoots had higher photosynthetic rates when PPFD was below 600 moles m^-2s^-1, and at 250 moles m^-2s^-1 shade shoots maintained about 50% of the net photosynthetic rate of sun shoots at light saturation. Shade shoots appeared capable of benefitting photosynthetically from elevated temperatures within krummholz mats despite relatively low light levels. Especially rapid photosynthesis may occur when canopy needles are illuminated by sunflecks and needle temperatures rise by 10° C or more.Snow cover appears crucial for the survival of needles during winter. Snow accumulated within krummholz needle canopies before the sub-canopy zone of unfoliated branches became filled. The concentrated needle growth in the krummholz canopy captured snow in early autumn without support from ground-level snowpack. Early snow cover in both species prevented cuticle abrasion and resulted in high winter needle water contents and viabilities for subsurface compared to surface needles which became abraded, severely dehydrated, and had high mortality between December and February, especially on windward sides of shoots.Extremely high concentrations of needles within krummholz mat canopies created an aerodynamic structure which elevated needle temperatures to more optimal photosynthetic levels in summer and resulted in more efficient snow accumulation in winter. These factors appear crucial for winter needle survival. Thus, krummholz mats appear to be an important adaptation in growth form which provides survival benefits in both summer and winter.     

Effects of ultraviolet radiation on early larval stages of the Alpine newt, Triturus alpestris, under natural and laboratory conditions

Although Alpine newts (Triturus alpestris) are found at altitudes up to 2500 m, their larvae proved to be extremely sensitive to UV radiation when exposed in clear tapwater to natural sunlight or to comparable artificial UV-B radiation in the laboratory. The experiments revealed severe skin damages (lysis of epithelial cells) and mortality after a few days of exposure. In their natural habitats above the timberline, however, the larvae are protected by the high concentration of dissolved organic carbon in the water leading to an almost complete absorption of UV radiation within the first few centimeters of the water layer. Furthermore, when exposed to UV radiation larvae show erratic swimming activities that may carry them into protected areas. Only in very shallow natural habitats did we detect sublethal UV-like histological effects. Shallow high mountain ponds with clear water normally lack newt populations, probably as a consequence of both low temperature and the effects of UV radiation. Received: 4 October 1996 / Accepted: 7 February 1997     

Winter-drought induced embolism in Norway spruce (Picea abies) at the Alpine timberline

At the timberline in the Central Alps, climatic conditions during winter frequently cause excessive drought stress (frost drought, 'Frosttrocknis'), which we hypothesized to induce cavitation in trees. We investigated the extent of winter-embolism in Norway spruce ( Picea abies (L.) Karst.) growing near the timberline and analysed adaptations in vulnerability and anatomy. We found conductivity losses of up to 100% at the highest elevation (2020 m) correlated with low water potentials down to − 4.0 MPa. Vulnerability thresholds (50% loss in conductivity) decreased from − 3.39 MPa at 800 m to − 3.88 MPa at 1600 m corresponding to a decrease in tracheid cross-sectional area as well as pit and pit pore diameters. These thresholds were lower than potentials measured in embolized twigs near the timberline at the sampling dates probably due to lower potentials and/or a role of freeze-thaw events earlier in winter. Data indicated refilling processes, which may be of particular relevance for trees at the timberline, since adaptations in drought-induced vulnerability failed to prevent winter-embolism.     

Sap flow rates of mangrove trees are not unusually low

 In contrast with previous reports, we observed high transpiration rates in mangrove trees. Maximum sap velocities and mean daytime sap flow rates were estimated from heat pulse velocity in entire, field grown trees of Avicennia cf. alba Blume and Rhizophora apiculata Blume. Results were within the range of values measured by identical techniques for trees in lowland dipterocarp and tropical heath forests with a similar climate in Brunei Darussalam (north Borneo). High stomatal conductance (400 mmol m^{–  2} s^{–  1}) was also measured for well insolated leaves of A. cf. alba, with midday water potentials reaching about  – 3 MPa in both species. Received: 11 September 1996 / Accepted: 27 January 1997     

Transpiration and drought resistance of Douglas-fir seedlings exposed to excess ammonium

 In a pot trial growth and transpiration of 3-year-old Douglas-fir seedlings on an acid, sandy soil was examined at a deficient (30 kg N ha^{–  1} year^{–  1}) and an excessive level (120 kg N ha^{–  1} year^{–  1}) of NH₄ application. Dissolved ammonium sulphate was applied to the pots weekly for two growing seasons. In half of the pots a complete set of other nutrients was applied in optimal proportions to the applied nitrogen. Water supply was optimal and transpiration was recorded. At the end of the second treatment season irrigation was stopped for 2 weeks during dry and sunny weather. Both high application of NH₄ and additional nutrients increased shoot growth and transpiration demand in the first treatment year. The root system was smaller at higher N level and this reduced water uptake accordingly. In the second year the combination of high NH₄ ⁺ and additional nutrients affected root functioning predominantly due to salinity effects and this seriously decreased water uptake capacity and shoot water potentials, finally resulting in tree death. Without addition of other nutrients the high NH₄ ⁺ application resulted in a high degree of soil acidification, which damaged the roots, that showed a decrease in water uptake capacity. At the low NH₄ supply level soil acidification was lower, and root functioning was not affected, and the trees recovered quickly from the imposed drought. Higher needle K and P status depressed transpiration rates at the low NH₄ application rate. Received: 9 January 1995 / Accepted: 18 September 1995     

Hydraulic efficiency and safety of leader shoots and twigs in Norway spruce growing at the alpine timberline

Xylem within trees varies in its hydraulic efficiency and safety. Trees at the alpine timberline were expected to exhibit a hydraulic architecture protecting the leader shoot from winter embolism. Hydraulic and related anatomical parameters were compared as well as seasonal courses of winter embolism in leader shoots and twigs of Norway spruce trees growing at 2000 m. Leader shoots had a 1.4-fold higher specific hydraulic conductivity (ks) as well as a 4.9-fold higher leaf specific conductivity (kl) than side twigs. Vulnerability to drought-induced embolism was lower in leader shoots with a 50% loss of conductivity occurring at a water potential (Psi 50) 0.7 MPa lower than in twigs. Higher ks and kl were related to 1.2-fold wider tracheid diameters in leader shoots. Lower vulnerability corresponded to smaller pit dimensions but not to wood density. High ks and kl reflect the hydraulic dominance of the leader shoot, which is important for its water supply during summer. Low vulnerability protects the leader shoot from embolism during the winter season. In field measurements at the timberline during the winter of 2001/2002, conductivity losses of up to 56% were observed only in twigs while leader shoots showed little or no embolism. Results demonstrate that leader shoot xylem is both hydraulically efficient and safe.     

Cortical microtubules rearrange during differentiation of vascular cambial derivatives, microfilaments do not

The plant cytoskeleton has been implicated in a variety of morphogenetic events in higher plants. Most of this work, however, has concentrated on epidermal cells or primary tissues. We have investigated the cortical microtubular (CMT) and microfilament (MF) components of the cytoskeleton in a secondary tissue  –  active vascular cambium of Aesculus hippocastanum L. (horse-chestnut)  –  and followed the changes in these components during the early stages of differentiation of fusiform cambial derivatives to axial elements of the secondary vascular system. A correlative approach was used employing indirect immunofluorescence microscopy of α-tubulin on 6 μm sections, and transmission electron microscopy of 60 nm sections. The study has demonstrated a rearrangement of the CMT cytoskeleton, from random to helical, as fusiform vascular cambial cells begin to differentiate as secondary phloem vascular tissue. A similar CMT rearrangement is seen as fusiform cambial cells begin to differentiate as secondary xylem fibres. This rearrangement is interpreted as evidence of determination of cambial derivatives towards vascular development. Axially-oriented MF bundles are present in fusiform cambial cells and their axial orientation is retained in the vascular derivatives at early stages of their development even though the CMTs have become rearranged. Received: 5 August 1996 /  Accepted: 23 September 1996     

Laurel forests in Tenerife,Canary Islands

 Stand structure and leaf area distribution of a laurel forest in the Agua García mountains of Tenerife are described. The site is situated at 820 m a.s.l., faces NNE, and has a humid mediterranean climate. Summer droughts are mitigated by relatively high air humidity and clouds. The natural mixed hardwood forest is composed of six major tree species: Laurus azorica (Seub.) Franco, Persea indica (L.) Spreng, Myrica faya Ait., Erica arborea L. and two species of Ilex (I. platyphylla Webb & Berth. and I. canariensis Poivet.). The experimental stand had a density of 1693 trees ha^– 1, a basal area of 33.7 m²ha^– 1, and a cumulated volume of above-ground parts of trees of 231 m³ ha^– 1 with a corresponding dry mass of 204 ton ha^– 1. Diameters at breast height ranged from 6 to 46 cm. Mean concentration of plant dry mass per volume was 1.17 kg m^– 3. The vertical pattern of leaf area distribution in individual trees for all tree species was characterized by a Gaussian-like curve. Stand leaf area index was 7.8. These evergreen, broad-leaved (laurisilva or lucidophyllous) forests represent a relic forest that was widespread in the Mediterranean region some 20 million years ago. Our data illustrate some of the structural characteristics of this historically widespread forest type. Received: 2 December 1994 / Accepted: 6 November 1995     

Tracing water uptake by jarrah (Eucalyptus marginata) trees using natural abundances of deuterium

 Seasonal change in the δ²H content of water from twig sap, soil, rainfall and groundwater were measured to determine the water sources accessed by jarrah (Eucalyptus marginata) trees at three sites in Western Australia with differing soils and depths to water table. During winter and spring the main contributor to the water uptake of the trees was stored water in the surface layers of the soil replenished by predominantly winter rainfall. With the onset of summer drought jarrah became more reliant on water from deeper down the profile. There was no clear evidence that jarrah could tap water from groundwater more than 14 m deep in deep sands. Defining the source of water for trees in deep lateritic soils using stable isotopes is hampered by the uniform deuterium profiles down most of the unsaturated zone and into the groundwater. There was a limited response in the δ²H values of sapwater in twigs to changes in the δ²H of the upper layers of the deep sand following input of rainfall in autumn. The damped response was related to the small variation in the δ²H composition of rainfall in most events during the year and the mixing in the tree of water extracted from different locations in the soil profile. Received: 21 August 1995 / Accepted: 3 December 1995     

Assimilation,allocation and utilization of carbon by 3-year-old Scots pine (Pinus sylvestris L .) trees during winter and early spring

 The photosynthetic capacity of frost-hardy and frost-sensitive needles of 3-year-old Scots pines and the allocation and utilization of assimilated carbon was examined during winter and early spring. The photosynthates of the whole trees were labelled by ¹⁴CO₂ fixation and after chase periods of from 7 days to 4 months under natural climatic conditions, the distribution of radiocarbon in the various tissues of the trees was determined. During winter maximal photosynthetic rates of 1-year-old needles were considerably lower than in summer when calculated on a leaf area basis. However, when related to the chlorophyll content these discrepancies disappeared. The decrease of the photosynthetic capacity upon frost-hardening could be attributed to a two- to three-fold reduction in the chlorophyll content of the needles. The pulse-chase experiments showed that photosynthesis during the cold season preferentially provides substrates for respiration. Half of the assimilated ¹⁴C was respired during the first week, and after chase periods of 3 – 4 months the trees contained not more than 10 – 20% of the radiocarbon. The carbon, which was exported by the needles, was translocated basipetally via the twigs and the stem to the roots. Whereas in the axial system incorporation of radiocarbon into storage compounds, like starch, and into cell wall material was almost negligible during the cold season, in the roots one-third of the radiocarbon was recovered from starch 2 months after the ¹⁴C-pulse. In contrast to the above-ground parts of the trees, where starch content was very low during winter, in the roots considerable amounts of starch, up to 450 μmol hexose units · g^– 1 DW, were found even during mid-winter. In early spring the radiocarbon in the cell wall-, lipid-, and starch-fraction accounted for more than 80% of the ¹⁴C recovered at that time from the axial system. Incorporation of minor quantities into the cell wall fraction of the roots during winter and early spring indicate continuous root growth during the cold period as well as in early spring. Whereas during winter the buds did not attract freshly assimilated carbon, in spring just before bud break substantial amounts of carbon were translocated from the needles into the buds. In contrast, remobilization of carbon, which had been assimilated during autumn of the previous year, and import into the sprouting buds could not be demonstrated. Received: 3 November 1995 / Accepted: 1 March 1996     

DNA sequence analysis of the C4 antigen WH: evidence for two mechanisms of expression

 Amino acid and protein analyses have allowed the construction of a model for the C4-based Rodgers and Chido blood group antigens. The single low-frequency allele (WH) in this blood group system, however, has not been characterized at the molecular level. Two WH⁺ donors were studied by C4 agarose gel electrophoreses, immunoblot studies using monoclonal anti-Rg: 1 or anti-Ch: 1, serological phenotyping, polymerase chain reaction-restriction fragment length polymorphism of their C4 genes, and DNA sequencing of the WH allele. The first donor had the C4A1, A3 phenotype; the C4A1 carried Ch: 1, 3, 6 (thus exhibiting reversed antigenicity) and the C4A3 carried the WH antigen. The amino acid sequence of the WH allele was PCPVLD at positions 1101 – 1106, S at position 1157, and VDLL at positions 1188 – 1191. A second donor typed as C4A2, A4, B1 and was also WH⁺. Immunoblot analysis showed that a C4B1 protein expressed Rg: 1. Sequence analysis of the C4B genes showed the amino acids LSPVIH at positions 1101 – 1106, S at position 1157, and ADLR at positions 1188 – 1191. Thus, the WH antigen is a conformational epitope that can arise through different mechanisms on either a C4A or C4B gene. Received: 22 November 1995 / Revised: 19 February 1996     

The subalpine vegetation of Mt. Vysokaya, central Sikhote-Alin

The subalpine vegetation structure of Mt. Vysokaya, the Central Sikhote-Alin, is described. This vegetation consists mainly of subalpine spruce-fir forest, a complex of subalpine meadows, shrubs, groves of Betula lanata (B. ermanii s.l.), krummholz of Pinus pumila and alpine tundras. Significant disturbances in the vegetation structure were noted, especially in the forest-tundra ecotone accompanying a sharp reduction of the belts of Betula lanata and Pinus pumila. The altitudinal level of the upper timberline reaches 1600 m a.s.l. which is 250 m less than the expected altitude calculated by Kira's warmth index. An undergrowth of scattered trees of Picea and Betula are growing up to the mountain top. Based on these data and a review of the literature, we concluded that a catastrophic lowering of the timberline and devastation of the subalpine vegetation belt occurred several centuries ago, probably as result of fires.     

Analysis of anchor residues in a naturally processed HLA-DR53 ligand

 The peptide motif of the HLA-DR53 (DRB4^*0101) molecule, which is associated with autoimmune diseases including Vogt-Koyanagi-Harada’s syndrome, was determined by peptide binding assay using human L plastin p581 – 595 peptide and its substituted analogues. L plastin p581 – 595 peptide is one of the naturally processed peptides bound to HLA-DR9/DR53 (DRB1^*0901/DRB4^*0101) molecules. The binding affinity of each peptide to the HLA-DR53 molecule was measured by fluorescence intensity of biotinylated peptides to L cell transfectants expressing HLA-DR53 molecules, followed by treatment with avidin-fluorescence. Binding of biotinylated peptides to HLA-DR53 molecules was not inhibited by all single-alanine-substituted nonbiotinylated peptides, indicating that the replaced position was important for binding to the HLA-DR53 moleule. The inhibitory motif is considered to be an HLA-DR53-specific binding motif, composed of a positively charged residue (K) at position 1, a hydrophobic residue (I) at position 4, positively charged residue (R or K) at position 8 or 9, and another hydrophobic residue (I) at position 10. This predicted motif is different from the binding motifs of other HLA-DR molecules. Received: 29 April 1996 / Revised: 16 June 1996     

Dendroecological reconstruction and interpretation of larch budmoth (Zeiraphera diniana) outbreaks in two central alpine valleys of Switzerland from 1470 – 1990

 Outbreaks of the larch budmoth (LBM) (Zeiraphera diniana) recur cyclically approximately every 7 to 10 years in subalpine larch-cembran pine and montane to subalpine larch-Norway spruce forests of the relatively dry valleys of the European Alps. By dendroecologically analyzing increment cores from 570 host (European larch –Larix decidua) and non-host trees (cembran pine –Pinus cembra, Norway spruce –Picea abies) through the use of skeleton plots, at least 57 (59) outbreaks could be reconstructed in the optimum Upper Engadine Valley (suboptimum Goms Valley), Switzerland, during the time period 1503 (1472) to 1990. The average interval between initial years of successive outbreaks was 8.58 (8.95) years, SD 1.66 (2.13) years. Over the centuries spatial shifts of LBM activity between the two study areas occurred, probably due to climatic changes. Clear, site-specific differences in LBM attack could only be found in the suboptimum area where high-lying (>1800 m) and/or south-facing stands were infested most. LBM-afflicted trees proved to be unsuitable for climate reconstructions because the impact of the persistently recurring outbreaks on tree growth is dominant. In order to provide sufficient information for a detailed ecological interpretation of the course of an outbreak, latewood widths and/or densities have to be analyzed in addition to the ring-widths. Received: 11 February 1995 / Accepted: 19 July 1996     

The neutrophil as an information transmitter in tumor inhibition by a streptococcal biological response modifier,OK-432

 Effective treatment of a rat transplanted ascites tumor by i. p. injection of a streptococcal biological response modifier, OK-432, was abrogated by selective in vivo depletion of neutrophils by a monoclonal antibody, RP-3. The mechanisms by which neutrophils participate in the therapeutic action of OK-432 were studied with Winn’s assay using peritoneal exudate cells periodically obtained from rats i. p. injected with this biological response modifier. Intraperitoneal resident macrophages were first activated with OK-432, and within 3 h, tumor-inhibitory activity had moved to the early exuded neutrophils. However, 6 h after injection, exuded macrophages were the only cells involved in tumor inhibition. Considered together with other findings, it is likely that, in this system, neutrophils may transmit information from resident macrophages to exuded inflammatory macrophages in a series of responses induced by i. p. injection of OK-432. Received: 29 April 1996 / Accepted: 27 July 1996