Trigonelline in mature seeds and developing seedlings of <Emphasis Type="Italic">Glycine max</Emphasis>

Trigonelline (TRG) is known as a compatible solute in response to stress as well as a cell cycle regulator, and is more concentrated in legumes than other non-legume dicots. Four Glycine max L. genotypes (Essex, ExF 67, Forrest and Stressland) were used to examine TRG concentration in seeds and seedlings exposed to 30 or 100 mM NaCl, and to determine the association of TRG concentrations in seedlings with seedling growth. Seed germination across genotypes was inhibited by elevated salinity (71–91 %) in ExF 67 and Forrest and by accelerated aging (77–92 %) in Forrest. Length of seedlings in most genotypes stressed with NaCl apparently decreased. The TRG content in mature seeds of four genotypes was 44.4–74.6 μg g⁻¹(d.m.). TRG content significantly increased during early young seedling development, but remained or significantly reduced in some genotypes stressed with NaCl.     

Relationship of Heliothis zea predators, parasitoids and entomopathogens to canopy development in soybean as affected by Heterodera glycines and weeds

The influence of canopy development in soybean on the survival of corn earworm, Heliothis zea (Boddie) (Lepidoptera: Noctuidae), egg and larval stages and population dynamics of arthropod fauna were evaluated in field trials during 1986–88 in eastern North Carolina. Soybean canopy size decreased as soybean cyst nematode, Heterodera glycines Ichinohe (Nematoda: Heteroderidae), initial population densities increased. Plant species composition of the soybean canopy was affected by weed population densities. Mortality of H. zea larvae due to parasitism and infection with entomopathogens was greater in closed canopy and (or) weedy soybeans than in very open and (or) weed free soybeans. Predation and parasitism of corn earworm eggs were similar across nematode and weed density treatments. Natural enemy populations increased to highest levels during July in closed canopy and (or) weedy soybeans, coinciding with availability of largest prey population reservoirs. A delay in colonization of very open and (or) weed free soybeans by beneficial arthropods until mid to late August allowed greater H. zea larval survival than in closed canopy and (or) weedy soybeans. Arthropod species richness was generally greatest in closed canopy and (or) weedy soybeans during mid to late July, with differences becoming nonsignificant in August and early September. Mean and maximum ambient temperatures were higher and relative humidities lower in open canopy than in closed canopy plots. These conditions were less favorable for development of pathogens and natural enemies.     

Comparison of Salt-Induced Osmotic Adjustment and Trigonelline Accumulation in Two Soybean Cultivars

21-d-old seedlings of the soybean (Glycine max) cvs. Essex and Forrest were treated with NaCl in a step-wise manner over 9 d (3 d 30 mM, 3 d 70 mM, and 3 d 100 mM) and maintained under 100 mM NaCl for an additional 14 d. During salt treatment, osmotic potential decreased more in cv. Forrest relative to cv. Essex. In non-stressed leaf tissue, cv. Forrest contained more trigonelline (TRG) relative to cv. Essex. During salt treatment, TRG amounts increased in cv. Forrest but were unchanged in cv. Essex. Both cvs. osmotically adjusted in response to salt stress; the maximal osmotic adjustment was 0.80 and 0.18 MPa in cv. Forrest and cv. Essex, respectively. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effect of Soil Temperature and pH on Resistance of Soybean to Heterodera glycines

Soybean cyst nematode (SCN), Heterodera glycines Ichinohe, is a major pest of soybean, Glycine max L. Merr. Soybean cultivars resistant to SCN are commonly grown in nematode-infested fields. The objective of this study was to examine the stability of SCN resistance in soybean genotypes at different soil temperatures and pH levels. Reactions of five SCN-resistant genotypes, Peking, Plant Introduction (PI) 88788, Custer, Bedford, and Forrest, to SCN races 3, 5, and 14 were studied at 20, 26, and 32 C, and at soil pH''s 5.5, 6.5, and 7.5. Soybean cultivar Essex was included as a susceptible check. Temperature, SCN race, soybean genotype, and their interactions significantly affected SCN reproduction. The effect of temperature on reproduction was quadratic with the three races producing significantly greater numbers of cysts at 26 C; however, reproduction on resistant genotypes remained at a low level. Higher numbers of females matured at the soil pH levels of 6.5 and 7.5 than at pH 5.5. Across the ranges of temperature and soil pH studied, resistance to SCN in the soybean genotypes remained stable.     

An Alternative Method for Evaluating Soybean Tolerance to Heterodera glycines in Field Plots

Alternate planting dates and periodic destruction of the previous year''s soybean crop as well as 1-year bare fallow were used to establish a range of population densities ofHeterodera glycines for the subsequent year. Soybean cultivar Coker 156 (susceptible, moderately tolerant) was compared to cultivars Essex (susceptible, intolerant) and Bedford (resistant) to evaluate tolerance at different H. glycines population densities established through the previous year''s treatments. Yield of Coker 156 was consistently intermediate between yields of Bedford and Essex in 1986 and 1987. Yield of Essex was negatively correlated (P = 0.05) with preplant egg numbers of H. glycines in 1987, whereas yield of Bedford and Coker 156 were not related to nematode density. Reproduction of H. glycines was greater (P = 0.05) on the moderately tolerant Coker 156 than on either of the other cultivars.     

Changes in the Heterodera glycines Female Index as Affected by Ten-year Cropping Sequences

The objective of this experiment was to measure the change in female index (FI) of Heterodera glycines from bioassays on Bedford, Peking, PI 89772, and PI 90763 soybean (Glycine max) for 12 cropping sequence treatments over a 10-year period. Cropping sequences included continuous plantings of Forrest, Peking, and D72-8927 soybean (all resistant to race 3); Bedford, Nathan, and D75-10710 soybean (all resistant to races 3 and 14); a Bedford-corn (Zea mays) rotation; a rotation of Bedford, Essex (susceptible), and Forrest; and a 70:30 blend of Bedford and Forrest. The FI from bioassays with PI 89772 and PI 90763 decreased over time from 24.3 to 1.6 with treatments involving continuous Bedford, Nathan, and D75-10710 and the Bedford-corn rotation. The FI increased in bioassays using Bedford with treatments involving Bedford, Nathan, D75-10710, the Bedford-Forrest blend, and the two rotations. Results of this field experiment confirm greenhouse experiments in which reciprocal changes occur in FI on PI 89772 and PI 90673 compared with FI on Bedford.     

Damage and Reproductive Potentials of Pratylenchus brachyurus and P. penetrans on soybean

Damage and reproductive potentials of Pratylenchus brachyurus and P. penetrans on soybean, Glycine max, cvs. Essex, Forrest, and Lee 68, were determined in microplot tests. Cultivar Essex was generally tolerant to P. brachyurus. Yield of Forrest was suppressed linearly with increasing P_i''s in the sandy soil (r = -0.92) and loamy sand soil (r = -0.99). Low to moderate P_i''s in the sandy clay loam gave an increase in yields as compared to plants without nematodes. Yield was not affected by this nematode in muck. Lee 68 was very sensitive to P. penetrans in microplots. Yield vs. P_i was fitted by a quadratic model (r = 0.82) with yield decreasing sharply as P_i''s were increased. The reproduction of both species decreased with increases in P_i. Lee 68 was a good host for P. penetrans, whereas Essex and Forrest were fair to poor hosts for P. brachyurus.     

Response of Resistant Soybean Plant Introductions to Meloidogyne incognita in Field Microplots

The response of two soybean plant introductions, PI 96354 and PI 417444, highly resistant to Meloidogyne incognita, to increasing initial soil population densities (Pi) (0, 31, 125, and 500 eggs/100 cm³ soil) of M. incognita was studied in field microplots for 2 years. The plant introductions were compared to the cultivars Forrest, moderately resistant, and Bossier, susceptible to M. incognita. Averaged across years, the yield suppressions of Bossier, Forrest, PI 417444, and PI 96354 were 97, 12, 18, and < 1%, respectively, at the highest Pi when compared with uninfested control plots. Penetration of roots by second-stage juveniles (J2) increased linearly with increasing Pi at 14 days after planting. At the highest Pi, 62% fewer J2 were present in roots of PI 96354 than in roots of the other resistant genotypes. Soil population densities of M. incognita were lower on both plant introductions than on Forrest. At 75 and 140 days after planting, PI 96354 had the lowest number of J2 in the soil, with 49% and 56% fewer than Forrest at the highest Pi. The resistance genes in PI 96354 should be useful in a breeding program to improve the level of resistance to M. incognita in soybean cultivars.     

油菜类型和品种对外来入侵杂草小子虉草的替代控制作用

外来入侵植物小子虉草（Phalaris minor Retz.）是世界公认的冬季农田恶性杂草,掌握农作物对其替代控制作用具有重要的研究价值。前期研究表明,油菜是替代控制小子虉草的优良农作物,然而,目前尚不清楚油菜类型与品种对其控制能力的影响。为此选取与小子虉草同域发生的不同类型（白菜型油菜、芥菜型油菜和甘蓝型油菜）油菜品种各3种,通过田间小区实验和室内化感作用测定,对比研究其对小子虉草的生长、繁殖、表型以及化感作用的影响。田间实验显示：竞争方式（种内或种间竞争）和油菜类型对小子虉草的地上生物量、种子数、株高、分枝数、叶面积和比叶面积存在极显著（P=0.0001）影响;而油菜品种对小子虉草的地上生物量（P=0.6064）、种子数（P=0.3577）、株高（P=0.4279）、分枝数（P=0.6357）、叶面积（P=0.8839）和比叶面积（P=0.3424）均无显著影响。3种类型油菜对小子虉草生长、繁殖以及表型的影响存在明显差异,其中芥菜型油菜对小子虉草的上述指标的影响最强,而白菜型油菜的影响最弱。室内生物测定显示,油菜对小子虉草具有化感抑制作用,当供试油菜叶片水提液浓度为0.1 g/mL时,小子虉草种子的萌发和幼苗的株高、根长、生物量均被显著抑制;研究也表明不同类型油菜对小子虉草的化感作用显著不同,同等条件下,芥菜型油菜对小子虉草的化感抑制作用最强。综上所述,油菜类型对外来入侵小子虉草的控制作用存在显著差异,其中芥菜型油菜对植物小子虉草的替代控制作用明显优于白菜型油菜和甘蓝型油菜,而其强的化感抑草特性或许是其强控草能力的原因之一。另外,本研究也为进一步利用油菜替代控制入侵植物小子虉草提供了参考。     

葡萄糖对小球藻(Chloralla sp. HN08)光合作用和生长的影响

【目的】探讨葡萄糖作为外加碳源对热带海洋小球藻(Chloralla sp.HN08)生物质生产和脂、光合色素、碳水化合物及可溶性蛋白等细胞主要成份含量的影响。【方法】分析比较小球藻HN08在光合自养和兼养(添加10 g/L葡萄糖)2种营养方式下的生长速率、细胞密度、光合放氧速率、油脂相对含量,以及可溶性总糖、淀粉和可溶性蛋白的含量。【结果】结果表明,在光照条件下葡萄糖(10 g/L)能促进小球藻(Chloralla sp.HN08)生长,提高细胞终密度,而异养条件下藻细胞逐渐衰亡。兼养条件下,细胞相对生长速率及细胞终密度分别是自养条件下的6.8倍和1.3倍。兼养藻细胞中可溶性糖、淀粉、油脂含量显著高于(P0.05)光合自养细胞,然而可溶性蛋白质和光合色素含量显著低于(P0.05)光合自养细胞。添加葡萄糖的小球藻液的光饱和点和呼吸速率均高于光自养条件下的细胞,但2种培养条件下藻液的净光合速率无显著差异(P0.05)。【结论】光照条件下,添加葡萄糖可显著提高小球藻HN08相对生长速率和细胞终密度,促进油脂与淀粉的积累。     

Effect of Within-field Variation in Soil Texture on Heterodera glycines and Soybean Yield

The influence of soil texture on Soybean yield in the presence of Heterodera glycines was investigated by comparing yields of susceptible cultivars with a resistant cultivar for 2 years. Soybean yield was negatively correlated with increasing sand content (P = 0.05). Yields of susceptible cultivars were suppressed with increasing sand content. Final nematode population densities were lowest in plots with greatest sand content. Soybean infection by SCN, as determined by the number of cysts 30 days after planting, was not consistently related to soil texture over 2 years. Initial nematode population density was positively related to soybean yield the first year and negatively related to soybean yield the second, probably a result of greater yield suppression by H. glycines in plots with greater sand content.     

降水对鄂尔多斯高原克隆植物分布的影响

鄂尔多斯高原地处半干旱区到干旱区的过渡带。通过沿鄂尔多斯高原从东到西的天然降水梯度的样带调查,分析了克隆植物的分布特征与降水的关系。结果表明:克隆植物的物种数量、占物种总数的比例和重要值以及根状茎型克隆植物的物种数量均与降水量呈显著正相关。5种优势植物克氏针茅、本氏针茅、黑沙蒿、赖草和短花针茅均为克隆植物,它们的分布也受降水影响。克氏针茅的密度、本氏针茅的盖度和密度、黑沙蒿的高度、盖度和密度以及赖草的高度均与降水量呈显著正相关。然而,赖草的盖度以及短花针茅的高度、盖度和密度均与降水量成负相关。因此,克隆植物在鄂尔多斯高原植被中具有重要作用,而且其作用随着降水量的增加而加强。     

Attempt to Select a Cyst Nematode Population on Soybean Plant Introduction 437654

A method of selecting soybean cyst nematode (Heterodera glycines Ichinohe) on segregating soybean progeny was evaluated for developing a population capable of reproducing on PI 437654. Direct selection on PI 437654 was not possible, since no cysts developed on it. Cysts were selected for 12 nematode generations on F₃ and F₄ plants of Forrest x PI 437654. No cysts of the selected population were produced on PI 437654, but more males were produced on it by the selected population than by the base population. The number of cysts on Forrest and other soybean lines considered to have some of the same genes for resistance increased with selection as expected. The increase in number of males on these other lines with some of the same genes for resistance as Forrest was greater than anticipated, indicating that these lines may have some of the same genes as PI 437654.     

Trypanosoma evansi: Adenosine deaminase activity in the brain of infected rats

The study was undertaken to evaluate changes in the activity of adenosine deaminase (ADA) in brains of rats infected by Trypanosoma evansi. Each rat was intraperitoneally infected with 10⁶ trypomastigotes either suspended in fresh (group A; n = 13) and cryopreserved blood (group B; n = 13). Thirteen animals were used as control (group C). ADA activity was estimated in the cerebellum, cerebral cortex, striatum and hippocampus. No differences (P > 0.05) in ADA activity were observed in the cerebellum between infected and non-infected animals. Significant (P < 0.05) reductions in ADA activity occurred in cerebral cortex in acutely (day 4 post-infection; PI) and chronically (day 20 PI) infected rats. ADA activity was significantly (P < 0.05) decreased in the hippocampus in acutely infected rats, but significantly (P < 0.05) increased in the chronically infected rats. Significant (P < 0.05) reductions in ADA activity occurred in the striatum of chronically infected rats. Parasites could be found in peripheral blood and brain tissue through microscopic examination and PCR assay, respectively, in acutely and chronically infected rats. The reduction of ADA activity in the brain was associated with high levels of parasitemia and anemia in acute infections. Alterations in ADA activity of the brain in T. evansi-infected rats may have implications for pathogenesis of the disease.     

Expression patterns in soybean resistant to Phakopsora pachyrhizi reveal the importance of peroxidases and lipoxygenases

Soybean rust caused by Phakopsora pachyrhizi Sydow is a devastating foliar disease that has spread to most soybean growing regions throughout the world, including the USA. Four independent rust resistance genes, Rpp1–Rpp4, have been identified in soybean that recognize specific isolates of P. pachyrhizi. A suppressive subtraction hybridization (SSH) complementary DNA (cDNA) library was constructed from the soybean accession PI200492, which contains Rpp1, after inoculation with two different isolates of P. pachyrhizi that result in susceptible or immune reactions. Both forward and reverse SSH were performed using cDNA from messenger RNA pooled from 1, 6, 12, 24, and 48 h post-inoculation. A total of 1,728 SSH clones were sequenced and compared to sequences in GenBank for similarity. Microarray analyses were conducted on a custom 7883 soybean-cDNA clone array encompassing all of the soybean-rust SSH clones and expressed sequence tags from four other soybean cDNA libraries. Results of the microarray revealed 558 cDNA clones differentially expressed in the immune reaction. The majority of the upregulated cDNA clones fell into the functional category of defense. In particular, cDNA clones with similarity to peroxidases and lipoxygenases were prevalent. Downregulated cDNA clones included those with similarity to cell-wall-associated protein, such as extensins, proline-rich proteins, and xyloglucan endotransglycosylases. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Assessment of potential sources of error in nitrogen fixation measurements by the nitrogen-15 isotope dilution technique

Although the use of ¹⁵N fertilizers to measure nitrogen (N₂) fixed in crops has increased substantially in recent years, some methodological uncertainties still remain unresolved. The results obtained from a greenhouse study of soybean [Glycine max. (L.) Merrill] inoculated by six different methods have been examined for potential errors arising from incorporating ¹⁵N labelled fertilizer into soil to estimate N₂ fixed in pods or shoots or the whole plant at three growth stages (50% flowering, pod-initiation and physiological maturity) using as reference crops, an uninoculated soybean cultivar and a non-nodulating soybean isoline. At the first harvest when N₂ fixed was very low, the estimates of N₂ fixed by the two reference crops did not match. At this stage the uninoculated soybean estimated about four times as much N₂ fixed in the symbiotic soybean as that measured using the non-nodulating soybean. For the second and third harvests, there were substantial increases in N₂ fixed, and both the non-nodulating and uninoculated soybean were equally suitable as reference crops for assessing N₂ fixed in the symbiotic soybean. These results indicate how critical and difficult the choice of the reference crop could be at early harvests, or when N₂ fixed is low. Even though there were significant differences in ¹⁵N enrichments in different organs (generally nodules < pods < roots < shoots), the estimates of N₂ fixed in soybean plants obtained by excluding roots and nodules did not differ much from those based on the whole plant. Of the above-ground organs, % N₂ fixed in pods (containing seeds) was closest to that of the whole plant (similar at P<0.05 at physiological maturity). However, the total N₂ fixed in pods or shoots was substantially lower than that fixed by the whole plant (P<0.05), although that for the pods and enclosed seeds once again was closer to N₂ fixed in the whole plant than that in the shoots.     

入侵害虫悬铃木方翅网蝽越冬后期在豫北寄主树干上的分布研究

【目的】入侵害虫悬铃木方翅网蝽近年来严重威胁我国城市行道树二球悬铃木的健康,受害严重时造成树木死亡。在越冬后期对悬铃木树干上的悬铃木方翅网蝽成虫的越冬习性开展研究,能为冬季防治积累生物学研究基础。【方法】早春随机调查河南省新乡市行道树二球悬铃木树干栓皮层外层下方隐藏的悬铃木方翅网蝽成虫,比较阳侧和阴侧虫口密度。同时记载悬铃木树干0.00~0.50 m、0.51~1.00 m、1.01~1.50 m、1.51~2.00 m 4个不同高度区段范围和树干皮层外层皲裂程度在5%以下、5%~25%、25%~50%、50%~75%、75%~95%、95%以上6个等级条件下害虫的虫口密度。【结果】栖息在悬铃木树干树皮下方的悬铃木方翅网蝽在阴面更多,阴面和阳面的虫口密度差异达极其显著水平[(F=6.63)(F0.01(1,19)=1.0132),P=0.00]。对寄主树干4个不同高度区段的虫口密度比较后,发现在树干1.0~1.5 m区段间的虫口密度最大,平均11.5头;且虫口密度在悬铃木不同高度区段间存在极显著差异[(F=26.91)(F0.01(3,56)=4.1519),P=0.00]。悬铃木树皮的不同皲裂程度对悬铃木方翅网蝽虫口密度的影响达极其显著水平[(F=31.02)(F0.01(3,56)=4.1519),P=0.00],在树皮外皮50%~75%皲裂程度条件下的虫口密度最大,平均9.8头。利用双因素方差分析方法分析树干不同高度和树皮不同皲裂等级的互作对虫口密度的分布影响达显著水平[(F=2.46)(F0.05(9,56)=2.0519),P=0.0195]。【结论】华北地区害虫秋冬防治时,在寄主悬铃木树干的最好涂白时机是在害虫全部下树后的12月上中旬进行,涂白高度以树干1.0~2.0 m范围为宜。本研究对悬铃木方翅网蝽在越冬期的预防和治理有较为重要的参考价值。     

Neoseiulus Californicus (Acari: Phytoseiidae) as a Potential Control Agent of Tetranychus Urticae (Acari: Tetranychidae): Effect of Pest/Predator Ratio on Pest Abundance on Strawberry

Neoseiulus californicus (McGregor) is a promising agent for successful Tetranychus urticae Koch control through conservation techniques, in strawberry crops in La Plata (Buenos Aires, Argentina). In prey–predator interaction, initial relative densities have an important effect on system dynamics. The economic threshold level (ETL) used for this pest in the present study was 50 active mites per leaflet. In our laboratory experiments, initial T. urticae to N. californicus ratio had a significant effect on the population abundance of T. urticae at a 7-day period. When pest/predator ratio was 5/1 (at initial pest densities from 5 to 15 females/leaflet) the final number of active T. urticae/leaflet was significantly lower than the ETL, while at 20 females/leaflet this number did not differ from the ETL. At 7.5/1 ratio, the final number of active T. urticae/leaflet, at initial pest densities from 5 to 15 females/leaflet, reached the ETL without surpassing it. At 10/1 and 15/1 ratios, pest densities exceeded the ETL only at 15 initial T. urticae/leaflet. Most greenhouse and field observations were consistent with the predictions of a graphical model based on experimental results. This predator was very effective in limiting pest densities at a 7-day period and within the range of pest–predator ratios and absolute densities used in this study. Conservation of N. californicus promoting favorable pest/predator ratios may result in early control of T. urticae.     

阔叶丰花草对荔枝园杂草群落特征及物种多样性的影响

【目的】明确阔叶丰花草在荔枝园定植对杂草群落特征及物种多样性的影响,为阔叶丰花草在果园应用提供参考依据。【方法】通过调查广东茂名高州荔枝国家现代农业产业园的荔枝园中90个有、无阔叶丰花草样方中杂草群落组成、群落高度以及物种多样性,探究阔叶丰花草定植对荔枝园杂草群落结构及物种多样性的影响。【结果】阔叶丰花草在荔枝园定植后,杂草种类减少,由对照样方的42种减少到试验样方的30种;杂草群落结构显著改变,牛筋草、白花鬼针草等恶性杂草重要值降低,薇甘菊、香附子、火炭母等杂草消失;阔叶丰花草的重要值与杂草群落高度和物种多样性指数为显著性负相关(P0.001),且均呈显著三项式回归关系。阔叶花草在荔枝园定植后可建立结构较为单一,且低矮的杂草群落。【结论】阔叶丰花草有望成为荔枝园生草的有益草种。     

Tolerance of a field grown soybean cultivar to elevated ozone level is concurrent with higher leaflet ascorbic acid level, higher ascorbate-dehydroascorbate redox status, and long term photosynthetic productivity

We examined the characteristics of ascorbic acid (ASC) level, dehydroascorbate (DHA) level, and the ASC–DHA redox status in the leaflets of two soybean cultivars grown in a field environment and exposed to elevated ozone (O₃) levels. These two cultivars, one that preliminary evidence indicated to be O₃-tolerant (cv Essex), and one that was indicated to be O₃-sensitive (cv Forrest), were grown in open-top chambers during the summer of 1997. The plants were exposed daily to a controlled, moderately high O₃ level (≈58 nl l⁻¹ air) in the light, beginning at the seedling stage and continuing to bean maturity. Concurrently, control plants were exposed to carbon-filtered, ambient air containing a relatively low O₃ level (≈24 nl l⁻¹ air) during the same period. Elevated O₃ did not affect biomass per plant, mature leaf area accretion, or bean yield per plant of cv Essex. In contrast, elevated O₃ level decreased the biomass and bean yield per plant of cv Forrest by approximately 20%. Daily leaflet photosynthesis rate and stomatal conductance per unit area did not decrease in either cultivar as a result of prolonged O₃ exposure. A 10% lower mature leaflet area in O₃-treated cv Forrest plants contributed to an ultimate limitation in long-term photosynthetic productivity (vegetative and bean yield). Possible factors causing cv Essex to be more O₃ tolerant than cv Forrest were: 1) mature leaflets of control and O₃-treated cv Essex plants consistently maintained a higher daily ASC level than leaflets of cv Forrest plants, and 2) mature leaflets of cv Essex plants maintained a higher daily ASC–DHA redox status than leaflets of cv Forrest plants. This revised version was published online in June 2006 with corrections to the Cover Date.