Distribution of pelagic blue-green algae in the North Pacific Ocean

The distribution of pelagic blue-green algae, especially ofTrichodesmium thiebautii, was investigated on the basis of the collection of theHakuhō Maru Cruise KH-69-4 along 155°W (50°N-15°S) in the North Pacific Ocean from September to November 1969.

1. Five species were identified:Trichodesmium thiebautii (most predominant),T. erythraeum, Oscillatoria sp.,Katagnymene spiralis andRichelia intracellularis.
2. T. thiebautii was most abundant in the western North Pacific central water and abundant next to it in the equatorial water, but it did not occur in the subarctic water.
3. T. thiebautii was ubiquitously distributed in the lower layer of 100–200 m in the equatorial water, though not in a large quantity.
4. T. thiebautii inhabited only the water warmer than 20°C. In its main habitat, nitrate and nitrite were almost zero, but ammonia and phosphate were present. There was not found any correlation between its occurrence and the salinity.
5. Blue-green algae were generally thinly populated in the water rich in diatoms.

     

Seasonal fluctuations of cell density of cyanobacteria and other picophytoplankton in Iwanai Bay,Hokkaido, Japan

In Iwanai Bay, which is located on the Japan Sea coast in Hokkaido, seasonal fluctuations in cell densities of phycoerythrin-dominant cyanobacteria and chlorophyll-dominant, other picophytoplankton were examined. Cell densities of cyanobacteria and other picophytoplankton ranged from 1.4×10⁵ to 1.9×10⁸ and from 4.0×10⁴ to 4.3×10⁶ cells 1^–1, respectively. The cell densities of both groups tended to increase after spring diatom bloom with remarkable fluctuations from June to August. This tendency was caused by the vertical distributions of both groups. The maximum density layer shifted from 0–20 m depth in April–June to 20–50 m depth in August–October. Cyanobacteria were dominant in picophytoplankton community which accounted for 73–99% of the total cell density during the whole year. Present results show that the picophytoplankton community in Iwanai Bay was influenced by a single water mass system (Tsushima Warm Current) during the whole year.This is contribution No. 254 from the Research Institute of North Pacific Fisheries, Faculty of Fisheries, Hokkaido University.     

北太平洋西边界流研究综述

综述了近20年来国内外学者在研究北太平洋西边界流的平均结构及NEC分叉动力机制、NM K流系平均输运的分配及变化、NM K流系季节及年际变化规律及其与EN SO之间的关系、NM K流系在热带和亚热带水交换中的作用以及水团的平均分布特征等方面所取得的主要成果。通过分析,发现东亚季风、R ossby波和K e lv in波等是影响北太平洋西边界流的主要因素;而缺乏长期直接的海流观测资料是深入研究北太平洋西边界流遇到的最大障碍。     

厦门西侧海域微微型浮游植物的时空分布及其调控机制

分别于１９９７年１０月与１９９８年４月,研究了厦门西侧海域微微型浮游植物类群组成,时空分布及其与环境因子的关系。结果表明：厦门西侧涨域微微型浮游植物在类群组成和丰度分布上存在的着在的时空变动;在类群组成上,富含藻蓝素的蓝细菌（ＰＣ细胞）在大多则站占优势（秋、春平均分别为７８％、４１％）,而真核微微型浮游植物（ＥＵ细胞）在九龙江口占优势（秋季高达８０％～１００％）;在丰度分布上,ＰＣ和富含藻红素的蓝细菌（ＰＥ细胞）呈秋季大于春季,而ＥＵ细胞秋春季则相近。与环境因子的回归分析表明,盐度是影响厦门西侧海域微微型浮游植物类群组成和丰度的关键因子,但其与细胞丰度的相关关系因季节不同而异;结果还表明,厦门西海域微微型浮游植物类群组成的多样性与该海域的异质性有关。     

印度洋海温异常对西北太平洋台风的影响及机制研究

基于近40 a NCEP/NCAR再分析月平均高度场、风场、涡度场、垂直速度场以及NOAA重构的海面温度(sea surface temperature,SST)资料和美国联合台风预警中心(Joint Typhoon Warning Center,JTWC)热带气旋最佳路径资料,利用合成分析方法,研究了前期春季及同期夏季印度洋海面温度同夏季西北太平洋台风活动的关系。结果表明:1)前期春季印度洋海温异常(sea surface temperature anomaly,SSTA)尤其是关键区位于赤道偏北印度洋和西南印度洋地区对西北太平洋台风活动具有显著的影响,春季印度洋海温异常偏暖年,后期夏季,110°～180°E的经向垂直环流表现为异常下沉气流,对应风场的低层低频风辐散、高层辐合的形势,这种环流形势使得低层水汽无法向上输送,对流层中层水汽异常偏少,纬向风垂直切变偏大,从而夏季西北太平洋台风频数偏少、强度偏弱,而异常偏冷年份则正好相反。2)春季印度洋异常暖年,西北太平洋副热带高压加强、西伸;而春季印度洋异常冷年,后期夏季西北太平洋副热带高压减弱、东退,这可能是引起夏季西北太平洋台风变化的另一原因。     

西太平洋冬季上层水体有色溶解有机物的分布和转化特征

为深入解析西太平洋溶解有机碳的生物地球化学过程,本研究于2015年12月至2016年1月,开展了西太平洋上层水体有色溶解有机物（CDOM）吸收光谱和荧光光谱特征研究。研究结果表明,西太平洋上层水体CDOM吸收系数a（320）变化范围为0.01~1.07 m-1,平均值为0.18 m-1;其较高值位于100~200 m水层,表层的海水相对含量较低,主要以有机物的光化学分解为主。采用PARAFAC分析CDOM三维荧光光谱特征,得到1种类腐殖质组分C2（252（310 nm）/405 nm）及2种类蛋白组分C1（224（276 nm）/335 nm）和C3（224（260 nm）/300 nm）,其中类腐殖质荧光组分占总荧光强度的11%~22%,蛋白质荧光组分占总荧光强度的78%~89%,蛋白质荧光中类色氨酸和类络氨酸组分对荧光强度的贡献相当。洋流在大尺度上控制西太平洋CDOM的分布特征,两流交界处和环流形成区域的CDOM相对含量较高,荧光信号较强。西太上层水体CDOM相对含量和荧光信息,与温度、盐度、DO和营养盐等理化因素之间的相关分析结果表明,CDOM主要成分类蛋白质的产生主要受上层水体初级生产过程控制。     

Methane in the western North Pacific

The concentration of methane in about 400 seawater samples collected in the western North Pacific, mostly from 40°N to 5°S along 165°E was determined. While the concentration of methane in the surface water was slightly greater in the high-latitudes, it did not widely vary with a standard deviation of 0.29 n mol/l for a mean value of 2.49 n mol/l. The 90% confidence limit of the mean was 0.08 n mol/l. The degree of oversaturation in 1991 (31±4%) was not different from that in circa 1970. If we assume that this degree of oversaturation occurs in the entire oceans, the annual flux of methane becomes 6×10¹²g CH₄. Both the concentrations of methane and chlorophylla were higher in the surface 100 m layer. However, the correlation between them was not well in the entire surface waters. This may indicate that the production of methane is not directly related to the photosynthetic process. The concentration of methane decreased gradually with increasing depth down to 1000 m. Its horizontally and vertically uniform concentration in the abyssal water suggests that the turnover time of methane in the oxic pelagic water is in the range between a few years and a few hundred years.     

Silicon flux and distribution of biogenic silica in deep-sea sediments in the western North Pacific Ocean

We investigated biogenic silica, several biological components, and silicate in pore-water in the abyssal sediment to determine silicon flux of western North Pacific during several cruises. The surficial sediment biogenic silica content was high at high latitudes with the boundary running along the Kuroshio Extension, and maximum values (exceeding 20%) were found in the Oyashio region. In the subtropical region to the south, most stations showed less than 5% biogenic silica content. This distribution pattern reflected primary production and ocean currents in the surface layer very well. Pore-water samples were collected from 4 stations along the east coast of Japan. The highest asymptotic silicic acid concentration (670 μmol L^?1) in pore-water was observed at the junction of Kuroshio and Oyashio, followed by samples from the Oyashio region. It is at the southern station that the lowest value (450 μmol L^?1) was observed, and the primary production is low under the influence of Kuroshio there. The diffusive flux followed the same geographic trend as the asymptotic silicic acid concentrations did, ranging 77–389 mmol m^?2 yr ^?1. Multiple sampling of pore-water was conducted throughout the year at one station at high latitude. The average annual biogenic silica rain flux observed using sediment traps was 373 mmol m^?2 yr^?1; the diffusive flux and burial flux at the sediment–water interface were 305 and 9 mmol m^?2 yr^?1, respectively. We concluded that most of the settling silica particles dissolved and diffused at the sediment–water interface and approximately 3% only were preserved in this area. In addition, the obvious time lag observed between the peak rain flux and the maximum diffusive flux suggested that primary production in the surface layer has a great influence on the sedimentation environment of abyssal western North Pacific. These transitions of Si flux at the sediment–water interface were considerably greater in northwestern North Pacific than in southwestern North Pacific. In addition, a station in the Philippine Sea indicated high biogenic silica content because of Ethmodiscus ooze, which are scattered randomly on the sea floor in the subtropical region.     

Dissolved and labile particulate Zr,Hf, Nb,Ta, Mo and W in the western North Pacific Ocean

Dissolved and labile particulate Zr, Hf, Nb, Ta, Mo and W were determined at stations K1 (51°N, 165°E), K2 (47°N, 160°E), KNOT (44°N, 155°E) and 35N (35°N, 160°E) in the western North Pacific Ocean. A portion of seawater for dissolved species (D) was passed through a 0.2 μm Nuclepore filter and acidified to pH 2.2 with HCl and HF. A portion of seawater for acid-dissolvable species (AD) was acidified without filtration. Labile particulate (LP) species is defined as AD minus D, which represents a chemically labile fraction of particulate species. D-Zr, Hf and Ta increase with depth, Nb shows a slight depletion in surface water, whereas Mo and W have a conservative vertical profile. The concentration range of D-Zr, Hf, Nb, Ta and W is 31–275, 0.14–0.95, 4.0–7.2, 0.08–0.29 and 40–51 pmol kg⁻¹, respectively, whereas that of Mo is 97–105 nmol kg⁻¹. LP-species of Zr, Hf and Ta account for 10–14% of AD in average and increase up to 25% below 4000 m, whereas those for Mo and W are negligible. In contrast, LP-Nb shows maxima (up to 27%) in surface water. We also found that D-Zr/Hf, Nb/Ta and Mo/W mole ratios generally increase in the order continental crust < river water < coastal sea < open ocean.     

Chlorofluorocarbons in the western North Pacific in 1993 and formation of North Pacific intermediate water

Chlorofluorocarbons (CFC-11 and CFC-12) in the intermediate water having between 26.4 and 27.2 were determined at 75 stations in the western North Pacific north of 20°N and west of 175.5°E in 1993. The intermediate water of 26.4–26.6 was almost saturated with respect to the present atmospheric CFC-11 in the zone between 35 and 45°N around the subarctic front. Furthermore, the ratios of CFC-11/CFC-12 of the water were also of those formed after 1975. These suggest that the upper intermediate water (26.4–26.6) was recently formed by cooling and sinking of the surface water not by mixing with old waters. The water below the isopycnal surface of 26.8 contained less CFCs and the area containing higher CFCs around the subarctic front was greatly reduced. However, the CFC age of the lower intermediate water (26.8–27.2) in the zone around the subarctic front was not old, suggesting that the water was formed by diapycnal mixing of the water ventilated with the atmosphere with old waters not containing appreciable CFCs, probably the Pacific Deep Water. The southward spreading rate decreased with depth and it was one sixth of its eastward spreading rate of the North Pacific Intermediate Water (NPIW).     

Bathymetric patterns of meiofaunal abundance and biomass associated with the Kuril and Ryukyu trenches, western North Pacific Ocean

The abundance and biomass of metazoan meiofauna and their relationships with environmental factors [chloroplastic pigment equivalents (CPE) and sediment characteristics] were studied quantitatively around and within the Kuril Trench (560-7090 m) and the Ryukyu Trench (1290-7150 m), which are located in eutrophic and oligotrophic regions, respectively, of the western North Pacific. Faunal abundance and biomass, as well as the CPE content of sediments, were considerably higher in the Kuril region than in the Ryukyu region. In both cases, CPE tended to decrease with water depth, but relatively high values were found in the deepest areas, suggesting that organic matter has accumulated in both trenches. Meiofaunal abundance and biomass were lower than expected from sediment CPE values at hadal stations below 6000 m. Differences in the density and biomass of meiofauna between these two trenches appeared to reflect differences in overall ocean productivity above them. When the analysis was restricted to each region, however, no association was found between the abundance and biomass of meiofauna and food availability. Furthermore, the factors regulating the bathymetric patterns in these meiofaunal parameters appeared to differ between the two trenches.     

Nutrient gradients in the western North Atlantic Ocean: Relationship to microbial community structure and comparison to patterns in the Pacific Ocean

Studies of nitrogen and phosphorus dynamics in the oligotrophic surface waters of the western North Atlantic Ocean have been constrained because ambient concentrations are typically at or below the detection limits of standard colorometric methods, except during periods of deep vertical mixing. Here we report the application of high-sensitivity analytical methods—determinations of nitrate plus nitrite (N+N) by chemiluminescence and soluble reactive phosphorus (SRP) by the magnesium induced co-precipitation (MAGIC) protocol—to surface waters along a transect from the Sargasso Sea at 26°N through the Gulf Stream at 37°N, including sampling at the JGOFS Bermuda Atlantic Time-series Study (BATS) station. The results were compared with data from the BATS program, and the HOT station in the Pacific Ocean, permitting cross-ecosystem comparisons. Microbial populations were analyzed along the transect, and an attempt was made to interpret their distributions in the context of the measured nutrient concentrations.Surface concentrations of N+N and SRP during the March 1998 transect separated into 3 distinct regions, with the boundaries corresponding roughly to the locations of the BATS station (∼31°N) and the Gulf Stream (∼37°N). Although N+N and SRP co-varied, the [N+N] : [SRP] molar ratios increased systematically from ∼1 to 10 in the southern segment, remained relatively constant at ∼40–50 between 31°N and 37°N, then decreased again systematically to ratios <10 north of the Gulf Stream. Dissolved organic N (DON) and P (DOP) dominated (⩾90%) the total dissolved N (TDN) and P (TDP) pools except in the northern portion of the transect. The [DON] : [DOP] molar ratios were relatively invariant (∼30–60) across the entire transect.Heterotrophic prokaryotes (operationally defined as “bacteria”), Prochlorococcus, Synechococcus, ultra- and nanophytoplankton, cryptophytes, and coccolithophores were enumerated by flow cytometry. The abundance of bacteria was well correlated with the concentration of SRP, and that of the ultra- and nanophytoplankton was well correlated with the concentration of N+N. The only group whose concentration was correlated with temperature was Prochlorococcus, and its abundance was unrelated to the concentrations of nutrients measured at the surface.We combined our transect results with time-series measurements from the BATS site and data from select depth profiles, and contrasted these North Atlantic data sets with time-series of N and P nutrient measurements from a station in the North Pacific subtropical gyre near Hawaii [Hawaii Ocean Time-series (HOT) site]. Two prominent differences are readily observed from this comparison. The [N+N] : [SRP] molar ratios are much less than 16 : 1 during stratified periods in surface waters at the BATS site, as is the case at the HOT site year round. However, following deep winter mixing, this ratio is much higher than 16 : 1 at BATS. Also, SRP concentrations in the upper 100 m at BATS fall in the range 1–10 nM during stratified periods, which is at least one order of magnitude lower than at the HOT site. That two ecosystems with comparable rates of primary and export production would differ so dramatically in their nutrient dynamics is intriguing, and highlights the need for detailed cross ecosystem comparisons.     

Distribution of particulate organic carbon and nitrogen in the Bering Sea and northern North Pacific Ocean

Particulate matter was collected in the Bering Sea and the northern North Pacific Ocean during the cruise of R. V. Hakuho-maru, Ocean Research Institute of Tokyo University in summer of 1975. The particulate matter was analyzed for organic carbon and nitrogen, chlorophylla and amino acids.The concentrations of particulate organic carbon and nitrogen were measured with the range of 16–422gC l^–1 and 1–85gN l^–1, 19–186gC l^–1 and 1–26gN l^–1, 46–1,038gC l^–1 and 6–79gN l^–1 and 19–246gC l^–1 and 2–25gN l^–1 in the Oyashio, the Deep Bering Sea, the continental shelf of Bering Sea and the northern North Pacific, respectively. Particulate organic carbon and nitrogen decreased with depth throughout the areas. The average concentrations of organic carbon and nitrogen in the entire water column tended to decrease in the following order; the continental shelf > Oyashio > northern North Pacific > Deep Bering Sea.C/N of particulate matter varied in the range of 3–15 (7 on average) in surface waters throughout the areas and these values tended to increase with depth to 5–20 (11 on average) in deep waters without significant regional variability.Linear regressions between chlorophylla and particulate organic carbon in the euphotic layers indicate that detrital organic carbon accounted for 34.2, 44.9, 49.1 and 25.2 % of particulate organic carbon in the Oyashio, the Deep Bering Sea, the continental shelf and the northern North Pacific, respectively.Particulate amino acid was determined in the range of 10.3–78.0g l^–1, 104–156g l^–1 and 10.4–96.4g l^–1 in the Deep Bering Sea, the continental shelf and the northern North Pacific, respectively. Aspartic acid, glutamic acid, serine, glycine and alanine were found as dominant species of amino acid of particulate matter.     

西太平洋秋季130°E断面有色溶解有机物的分布特征及光降解行为研究

对2018年秋季西太平洋130°E断面上层水体有色溶解有机物(CDOM)的光学特性及光降解行为进行了研究。结果表明,西太平洋上层水体CDOM的吸收系数a(320)变化范围为0.025～0.64 m^-1,平均值为(0.20±0.08) m^-1;a(320)在表层相对较低,主要与表层CDOM的光漂白去除有关;在100～200 m水层较高,主要与次表层的生物活动有关。利用三维荧光光谱-平行因子分析技术,识别出两种荧光组分:类酪氨酸组分C1和海洋类腐殖质组分C2。C1主要源于棉兰老冷涡-上升流所带来的营养物质对浮游植物生产活动和微生物活动的促进作用;C2主要源于黑潮所带来的海洋类腐殖的输入。光化学降解实验发现,CDOM吸收值的损失主要发生在紫外波段;光照60 h后,类酪氨酸组分相较于海洋类腐殖质组分更易发生光降解;且光降解是西太平洋海域CDOM的重要去除途径。     

Bottom water temperature measurements in the South China Sea,eastern Indian Ocean and western Pacific Ocean

文章报道了一批新的海底底水温度(BWT)数据,其中南海(SCS)158个站位、东印度洋(EIO)30个站位及西太平洋(WPO)37个站位。基于这批新的BWT数据,获得南海和西太平洋海域底水温度与水深经验关系,可为地球物理和物理海洋提供准确、可靠的海底温度边界。这将有助于海底油气资源调查与评估。同时,这批实测数据表明:1)水深超过3500m的海域,其底水温度在南海约为2.47℃,比东印度洋(～1.34℃)和西太平洋(～1.60℃)稍微偏高。这与大洋传送带模式所预测的情况比较吻合。该模式认为:低温高盐的海水,从北大西洋格陵兰岛和冰岛附近海域下沉到深层,然后向南流动,再与南极洲周围海域的低温高盐海水一同向北进入印度洋和太平洋。而南海是一个相对比较封闭的热带边缘海,其内部海水与印度洋和菲律宾海交换有限,导致海水温度整体高于印度洋和太平洋。2)台西南盆地水深在2700～3000m的部分站位,其底水温高达约3.00℃,明显高于其周边同水深海域底水温度(平均值约为2.33℃)。这可能是台西南盆地海底水热活动导致的结果。3)在东印度洋和西太平洋水深超过4800m海域,底水温度随着水压增大稍有升高,其升高率分别为10.6mK·MPa~(-1)和12.0mK·MPa~(-1)。这与理论估算的深层底水绝热压力温度梯度范围较为吻合。这也意味着东印度洋和西太平洋深层底水,主要由绝热自压作用导致其温度随着深度的增大而升高。     

Abundance and size composition of the summer phytoplankton communities in the western North Pacific Ocean,the Bering Sea,and the Gulf of Alaska

Chlorophylla concentrations (Chla) of size-fractionated phytoplankton samples were measured in the western North Pacific Ocean, the Bering Sea, and the Gulf of Alaska during the summer of 1986. Among samples collected in the upper 100 m (total of 210 samples), 207 samples were dominated by micro- (>10 m) or picoplankton (<2 m) and only three samples were represented by nanoplankton (2–10 m). These 207 samples were classified based on the total Chla content into three types: Type H (>1.0 g l^–1), Type M (0.5–1.0 g l^–1), and Type L (<0.5 g l^–1). These types further divided into two subtypes (-p and-m), depending upon dominancy of pico (-p) and microplankton (-m). The phytoplankton community was represented by Type L-p in the Gulf of Alaska, where 80% of the samples fell into this type. It was represented by Type M-p in the western North Pacific and the Oceanic Domain in the Bering Sea, where 53 and 41% of samples were identified as this type, respectively. In the Middle Domain of the Bering Sea, 68% of samples collected below the nitracline was Type H-m, which indicates blooms of microplanton. This type was also observed in the neritic waters near the Aleutian Islands. These types described above are consistent with a general trend that an increase in phytoplankton abundance is attributed to the growth of microplankton. An unusual type occurred above the nitracline of the Middle Domain, where microplankton prevailed, although the total Chla was less (Type L-m). This type represents a feature of late phase of an ice edge bloom. Another unusual type was found mainly in the Outer Domain of the Bering Sea, where the total Chla was high and picoplankton prevailed (Type H-p). The predominance of picoplankton seems to result from the heavy grazing intensity of large calanoid copepods upon microplankton but not upon picoplankton     

热带北太平洋西部固氮速率及固氮微生物组成

In the present study, we report N_2 fixation rate(~(15)N isotope tracer assay) and the diazotroph community structure(using the molecular method) in the western tropical North Pacific Ocean(WTNP)(13°–20°N, 120°–160°E). Our independent evidence on the basis of both in situ N_2 fixation activity and diazotroph community structure showed the dominance of unicellular N_2 fixation over majority of the WTNP surface waters during the sampling periods.Moreover, a shift in the diazotrophic composition from unicellular cyanobacteria group B-dominated to Trichodesmium spp.-dominated toward the western boundary current(Kuroshio) was also observed in 2013. We hypothesize that nutrient availability may have played a major role in regulating the biogeography of N_2 fixation.In surface waters, volumetric N_2 fixation rate(calculated by nitrogen) ranged between 0.6 and 2.6 nmol/(L·d) and averaged(1.2±0.5) nmol/(L·d), with 10 μm size fraction contributed predominantly(88%±6%) to the total rate between 135°E and 160°E. Depth-integrated N_2 fixation rate over the upper 200 m ranged between 150 μmol/(m~2·d)and 480 μmol/(m~2·d)(average(225±105) μmol/(m~2·d). N_2 fixation can account for 6.2%±3.7% of the depthintegrated primary production, suggesting that N_2 fixation is a significant N source sustaining new and export production in the WTNP. The role of N_2 fixation in biogeochemical cycling in this climate change-vulnerable region calls for further investigations.     

Decadal-Scale Climate and Ecosystem Interactions in the North Pacific Ocean

Decadal-scale climate variations in the Pacific Ocean wield a strong influence on the oceanic ecosystem. Two dominant patterns of large-scale SST variability and one dominant pattern of large-scale thermocline variability can be explained as a forced oceanic response to large-scale changes in the Aleutian Low. The physical mechanisms that generate this decadal variability are still unclear, but stochastic atmospheric forcing of the ocean combined with atmospheric teleconnections from the tropics to the midlatitudes and some weak ocean-atmosphere feedbacks processes are the most plausible explanation. These observed physical variations organize the oceanic ecosystem response through large-scale basin-wide forcings that exert distinct local influences through many different processes. The regional ecosystem impacts of these local processes are discussed for the Tropical Pacific, the Central North Pacific, the Kuroshio-Oyashio Extension, the Bering Sea, the Gulf of Alaska, and the California Current System regions in the context of the observed decadal climate variability. The physical ocean-atmosphere system and the oceanic ecosystem interact through many different processes. These include physical forcing of the ecosystem by changes in solar fluxes, ocean temperature, horizontal current advection, vertical mixing and upwelling, freshwater fluxes, and sea ice. These also include oceanic ecosystem forcing of the climate by attenuation of solar energy by phytoplankton absorption and atmospheric aerosol production by phytoplankton DMS fluxes. A more complete understanding of the complicated feedback processes controlling decadal variability, ocean ecosystems, and biogeochemical cycling requires a concerted and organized long-term observational and modeling effort. This revised version was published online in July 2006 with corrections to the Cover Date.