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Organic and inorganic nitrogen nutrition of western red cedar, western hemlock and salal in mineral N-limited cedar-hemlock forests.

Western red cedar (Thuja plicata Donn.), western hemlock (Tsuga heterophylla Raf. Sarge) and salal (Gaultheria shallon Pursh) are the main species growing in cedar-hemlock forests on Vancouver Island, Canada. Based on the dominance of organic N in these systems, we tested the hypotheses that: (1) organic N can be utilized by the three plant species; and (2) salal, which is ericoid mycorrhizal and has high tannin concentration in its tissues, would absorb more N from the complex organic N compounds than the other two species. The abilities of cedar, hemlock and salal to take up 15N,13C-labelled glutamic acid were measured and the capacities of the three species to use nitrate (NO3-), ammonium (NH4+), glutamic acid, protein and protein-tannin N were compared over a 20-day period. Based on 13C enrichment, all three species absorbed at least a portion of glutamic acid intact. Cedar, hemlock and salal also showed similar patterns of N uptake from the NO3-, NH4+, glutamic acid, protein and protein-tannin treatments. The largest proportions of applied N were taken up from the NO3- and NH4+ treatments while smaller amounts of N were absorbed from the organic N compounds. Thus organic N was accessed to a modest degree by all three species, and salal did not have a greater capacity to utilize protein and protein-tannin-N.

Carbon Isotopes↗

Volatile emissions of eastern hemlock, Tsuga canadensis, and the influence of hemlock woolly adelgid.

The volatile emissions of eastern hemlock, Tsuga canadensis Carriere, were identified and quantified using standard and chiral gas chromatography and mass spectrometry. All of the identified compounds were monoterpenes, and included alpha-pinene, myrcene, tricyclene, camphene, alpha-phellandrene, beta-pinene, limonene, beta-phellandrene, terpinolene, and bornyl acetate. alpha-Pinene, myrcene, and camphene comprised greater than 75% by mass of the total release. Infestation by the exotic insect, hemlock woolly adelgid (HWA, Adelges tsugae Annand), resulted in an increased release rate of monoterpenes from branch tips. Release rate was negatively correlated to the amount of the branch tip sample that was new growth, suggesting that release rate is greater from previous-year foliage. Additionally the percent composition of the volatile profile is slightly altered by infestation, with alpha-pinene comprising 57% of volatiles from infested foliage and 66% from uninfested foliage.

Animals↗

Comparison of numerical response and predation effects of two coccinellid species on hemlock woolly adelgid (Homoptera: Adelgidae).

The hemlock woolly adelgid, Adelges tsugae Annand, is an introduced pest in North America that is native to Asia, and is causing extensive damage to eastern hemlock (Tsuga canadensis Carriere) and Carolina hemlock (Tsuga caroliniana Englemann) in the eastern United States. We compared two coccinellids imported for biological control of the adelgid: Scymnus ningshanensis Yu et Yao from China and Pseudoscymnus tsugae Sasaji and McClure from Japan. In a laboratory study, we measured the numerical response of each beetle species to a range of prey densities, and in field studies we examined the reproductive success and ability of the coccinellids to reduce populations of the hemlock woolly adelgid. In the laboratory, S. ningshanensis showed a positive numerical response as hemlock woolly adelgid density increased, and P. tsugae showed a density-independent response. In field cages, the presence of S. ningshanensis resulted in negative hemlock woolly adelgid population growth, in contrast to positive growth in both control cages and cages containing P. tsugae. Both our laboratory and field experiments suggest that S. ningshanensis has good potential as a biological control agent of hemlock woolly adelgid.

Animals↗

The right-to-die movement: extrapolating from the National Hemlock Society U.S.A. membership survey.

A national membership survey of Hemlock Society USA was conducted by Fox and Kamakahi (1995). Respondents (N=6398) were asked a variety of questions, but in this paper we perform a longitudinal analysis of the characteristics of Hemlock Society USA members. Hemlock Society USA members are divided into three 5-year cohorts: Early Joiners (11 or more years of membership), Middle Joiners (6 to 10 years membership), and Late Joiners (5 or fewer years of membership). Differences between cohorts are examined and extrapolations made regarding Hemlock Society USA and the Right-to-Die Movement. A series of one-way ANOVAs were used with Scheme post-hoc comparisons as heuristic tools for assessing between-cohort differences. Late Joiners are different from earlier members, but are more like other Hemlock Society USA members than the adult U.S. population at large. Hemlock Society USA members are essentially societal "elites" (based on socio-demographic variables) who work in social environments that are decidedly split on the issue of voluntary suicide and euthanasia.

Euthanasia, Active, Voluntary↗

Vertical gradients in photosynthetic light response within an old-growth Douglas-fir and western hemlock canopy.

We examined needle-level light response of photosynthesis across a vertical light gradient within 45-55-m-tall western hemlock (Tsuga heterophylla (Raf.) Sarg.) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) trees growing in a 400-500-year-old mixed species stand. We determined: (1) whether light-saturated photosynthetic rates, light compensation points, and respiration rates varied from the upper to the lower canopy, and (2) if light-saturated photosynthetic rates, light compensation points, and respiration rates varied between Douglas-fir and western hemlock. Over a 25-m gradient from the canopy top to the lower canopy, mean light-saturated photosynthetic rates, light compensation points, and respiration rates declined in overstory Douglas-fir and western hemlock needles, paralleling a 65% decline in the mean daily photosynthetic photon flux density (PPFD). At the canopy top, increasing light-saturated photosynthetic rates relative to lower canopy needles increased carbon uptake at high PPFD. In the lower canopy, reduced respiration rates relative to upper canopy needles increased carbon uptake at low PPFD by reducing the light compensation point. At all canopy positions, western hemlock had lower mean light-saturated photosynthetic rates, light compensation points and respiration rates than Douglas-fir. As a result, western hemlock had higher net photosynthetic rates at low PPFD, but lower net photosynthetic rates at high PPFD compared with Douglas-fir.

Journal Article↗

Toxicoses in livestock from the hemlocks (Conium and Cicuta spp.).

The hemlocks, Conium maculatum (poison-hemlock) and Cicuta spp. (waterhemlock), are poisonous plants that cause sizeable losss to the livestock industry. Clinical signs of poisonhemlock toxicosis are similar in all species of livestock and include muscular weakness, incordination, trembling, initial central nervous system stimulation, depression and death from respiratory paralysis. Poison-hemlock also causes skeletal defects in the offspring of cattle, pigs and sheep and cleft palate in pigs when ingested during specific periods of gestation. The primary toxicants in poison-hemlock are coniine and gamma-coniceine. Coniine predominates in mature plants and seed, whereas gamma-coniceine predominates in early growth of the plant. Waterhemlock is the most violently toxic poisonous plant known. The toxicant is cicutoxin, which acts on the central nervous system, causing violent convulsions and death. Clinical signs of poisoning appear within 15 min after ingestion of a lethal dose and include excessive salivation, nervousness, tremors, muscular weakness and convulsive seizures interspersed by intermittent periods of relaxation and a final paralytic seizure resulting in anoxia and death. Elevated activities of lactic dehydrogenase, aspartate aminotransferase and creatine kinase in blood are observed, indicative of muscular damage. Toxicoses from poisonhemlock and waterhemlock generally occur in early spring when both plants emerge before other, more palatable plants begin to grow. All parts of the poison-hemlock plant are toxic. The root or tubers of waterhemlock are toxic; however, experimental evidence concerning the toxicity of other plant parts is inconclusive.

Alkaloids↗

Lupines, poison-hemlock and Nicotiana spp: toxicity and teratogenicity in livestock.

Many species of lupines contain quinolizidine or piperidine alkaloids known to be toxic or teratogenic to livestock. Poison-hemlock (Conium maculatum) and Nicotiana spp. including N. tabacum and N. glauca contain toxic and teratogenic piperidine alkaloids. The toxic and teratogenic effects from these plant species have distinct similarities including maternal muscular weakness and ataxia and fetal contracture-type skeletal defects and cleft palate. It is believed that the mechanism of action of the piperidine and quinolizidine alkaloid-induced teratogenesis is the same; however, there are some differences in incidence, susceptible gestational periods, and severity between livestock species. Wildlife species have also been poisoned after eating poison-hemlock but no terata have been reported. The most widespread problem for livestock producers in recent times has been lupine-induced "crooked calf disease." Crooked calf disease is characterized as skeletal contracture-type malformations and occasional cleft palate in calves after maternal ingestion of lupines containing the quinolizidine alkaloid anagyrine during gestation days 40-100. Similar malformations have been induced in cattle and goats with lupines containing the piperidine alkaloids ammodendrine, N-methyl ammodendrine, and N-acetyl hystrine and in cattle, sheep, goats, and pigs with poison-hemlock containing predominantly coniine or gamma-coniceine and N. glauca containing anabasine. Toxic and teratogenic effects have been linked to structural aspects of these alkaloids, and the mechanism of action is believed to be associated with an alkaloid-induced inhibition of fetal movement during specific gestational periods. This review presents a historical perspective, description and distribution of lupines, poison-hemlock and Nicotiana spp., toxic and teratogenic effects and management information to reduce losses.

Alkaloids↗

Genetic evaluation of alternative silvicultural systems in coastal montane forests: western hemlock and amabilis fir.

Genetic diversity and mating system were quantified for shelterwood, patch cut and green tree-retention silvicultural systems, and compared to adjacent old-growth. This is a component of a larger study conducted in montane old-growth forests of coastal British Columbia to evaluate the feasibility and ecological consequences of alternative silvicultural systems. The experiment includes replicated treatments representing a range of overstory removal adjacent to old-growth and clearcut areas. Based on 22 electrophoretically assayed loci, the effects of silvicultural systems on genetic parameters of amabilis fir (Abies amabilis and western hemlock (Tsuga heterophylla were assessed by comparing an average number of alleles per locus, the percent polymorphic loci, and observed and expected heterozygosity between parental populations and naturally regenerated progeny as well as among treatments. Genetic variation in natural regeneration was greater than in parental populations, especially for low-frequency alleles. Silvicultural treatments caused no significant differences in amabilis fir genetic-diversity parameters, while the shelterwood system resulted in lower observed and expected heterozygosity in western hemlock. Nei's genetic distance revealed that all parental populations were extremely similar. The two species had contrasting mating system dynamics with amabilis fir producing higher levels of correlated paternity and inbreeding with wider variation among individual tree outcrossing-rate estimates. Western hemlock had significant levels of correlated paternity only for the green tree and shelterwood treatments demonstrating family structuring inversely related to stand density. Inbreeding in western hemlock was significant but lower than that observed for amabilis fir with a J-shaped distribution for individual tree multilocus outcrossing-rate estimates. The pollination and dispersal mechanisms of the two species represent the most-likely factors causing these differences. Artificial regeneration may be utilized to augment the genetic resources of natural ingress.

Abies↗

A terrain-based paired-site sampling design to assess biodiversity losses from eastern hemlock decline.

Biodiversity surveys are often hampered by the inability to control extraneous sources of variability introduced into comparisons of populations across a heterogenous landscape. If not specifically accounted for a priori, this noise can weaken comparisons between sites, and can make it difficult to draw inferences about specific ecological processes. We developed a terrain-based, paired-site sampling design to analyze differences in aquatic biodiversity between streams draining eastern hemlock (Tsuga canadensis) forests, and those draining mixed hardwood forests in Delaware Water Gap National Recreation Area (USA). The goal of this design was to minimize variance due to terrain influences on stream communities, while representing the range of hemlock dominated stream environments present in the park. We used geographic information systems (GIS) and cluster analysis to define and partition hemlock dominated streams into terrain types based on topographic variables and stream order. We computed similarity of forest stands within terrain types and used this information to pair hemlock-dominated streams with hardwood counterparts prior to sampling. We evaluated the effectiveness of the design through power analysis and found that power to detect differences in aquatic invertebrate taxa richness was highest when sites were paired and terrain type was included as a factor in the analysis. Precision of the estimated difference in mean richness was nearly doubled using the terrain-based, paired site design in comparison to other evaluated designs. Use of this method allowed us to sample stream communities representative of park-wide forest conditions while effectively controlling for landscape variability.

Data Collection↗

Clinical spectrum of accidental hemlock poisoning: neurotoxic manifestations, rhabdomyolysis and acute tubular necrosis.

In the past, hemlock poisoning was only known for its neurotoxic effects; quite recently non-neurological features, consisting of rhabdomyolysis and acute renal failure, have been also described. Here we report our experience with these clinical findings, which we frequently observe in accidental hemlock poisoning. Between 1972 and 1990 we studied 18 patients: 17 of them were poisoned by conline (an alkaloid of Conium maculatim) in Apulia (Italy), and one by cicutoxin (the active principle of water hemlock) in New Mexico (USA). In the non-rapidly-fatal cases we tested myoglobinuria, serum muscle enzymes, and renal function. In the patients with acute renal failure we performed microscopical examination of kidney specimens; immunohistochemistry was carried out to identify myoglobin and actin in tubules. Coniine was detected in urine, serum, or tissues. Neurological features were present in all of our cases: coniine had a curare-like effect on the neuromuscular junction, whereas cicutoxin was convulsant on the central nervous system. In addition rhabdomyolysis was noted in the 17 subjects poisoned by coniine. Acute renal failure was observed in five patients; it was confirmed by histological evidence of tubular necrosis with intratubular deposition of myoglobin and actin released by rhabdomyolysis. Our cases seem to be the first with histopathologically proven acute tubular necrosis in coniine intoxication. In conclusion, in hemlock poisoning neurotoxic manifestations may be accompanied by rhabdomyolysis and acute tubular necrosis; increased awareness of these clinical features is recommended in order to improve the diagnostic and therapeutic procedure.

Adult↗

Successful transplantation of donor organs from a hemlock poisoning victim.

BACKGROUND: The poison hemlock plant (Conium maculatum) has been a known poison since early in human history, most notably as the agent used for the execution/suicide of Socrates in ancient Greece. No experience has been reported regarding the suitability of a hemlock victim's organs for transplantation. METHODS AND RESULTS: This report documents successful transplantation of the liver, kidney, and pancreas from a 14-year-old girl who died of anoxic encephalopathy from asphyxia after the accidental ingestion of fresh hemlock while on a nature hike. Predonation laboratory values were not remarkable, and liver and kidney biopsy results were normal. All organs in the three recipients had immediate function, and no recipient had any clinical evidence of transmitted toxin. All recipients are well, with functioning transplants at greater than 6 months after transplantation. CONCLUSIONS: Poison hemlock intoxication does not seem to be a contraindication to organ donation.

Adolescent↗

Water hemlock (Cicuta douglasii) toxicoses in sheep: pathologic description and prevention of lesions and death.

Water hemlock causes numerous livestock losses in North America every year. Description of pathologic and serum biochemical changes has been lacking in the literature. Tubers of western water hemlock (Cicuta douglasii) induced excessive salivation, tremors, grand mal seizures, skeletal and cardiac myodegeneration, and death in sheep given 1.2-2.7 g fresh tuber/kg body weight by gavage. Seizures were intermittent with periods of relaxation until death occurred from anoxia during seizure activity. In sheep given 1.5-2.5 times the lethal dose of water hemlock by gavage, intravenous administration of sodium pentobarbital at the onset of the first seizure prevented further seizure activity and skeletal and cardiac myodegeneration and resulted in rapid and complete recovery.

Animals↗

Carbon exchange of an old-growth eastern hemlock (Tsuga canadensis) forest in central New England.

Carbon (C) exchange of an approximately 200-year-old eastern hemlock (Tsuga canadensis L.) forest in central Massachusetts, USA, was estimated from mid-October 2000 through October 2001 based on eddy covariance measurements and statistical modeling from microclimatic data. Measurements were made in 68% of the hours during the year of study, with > 50% coverage in all months except December and August. Data were filtered by wind direction and atmospheric turbulence to remove invalid measurements. Analysis of filtered data showed that photosynthetically active radiation (PAR) was significant in predicting C exchange, except during the winter. Daily minimum air temperature affected C exchange in autumn and winter, whereas time of day, water vapor pressure deficit and air temperature had significant effects on C storage in spring, summer and fall. Most C storage in the stand occurred in April through July and in October 2001, with maximum rates in April and May. Persistent cold weather prevented C storage in December through March. In early spring 2001, C uptake was sensitive to nocturnal frost: daily minimum air temperatures below 0 degrees C reduced C fixation, and minima below -5 degrees C caused its virtual cessation. Soil temperature was a poor predictor of C balance during this period. In August, high soil and air temperatures (averaging 16.7 and 21.1 degrees C, respectively) drove high ecosystem respiration, which approximately balanced C uptake. These patterns show potential for stimulated C storage in hemlock forests in a warmer climate with fewer spring and autumn frosts, but reduced C storage during warmer summers. Estimated annual C storage was 3.0 Mg ha(-1), which is higher than for younger coniferous and deciduous forests during earlier years in the northeastern USA. Long-term data are needed to determine if the estimated high C storage in this hemlock forest is a result of interannual climate variation or an effect of forest composition.

Biomass↗

Feeding preference of three lady beetle predators of the hemlock woolly adelgid (Homoptera: Adelgidae).

In a laboratory study, we tested the feeding preferences of three coccinellid predators of hemlock woolly adelgid, Adelges tsugae Annand, an introduced pest of hemlock in the eastern United States. The species tested were Sasajiscymnus tsugae Sasaji & McClure (formerly Pseudoscymnus tsugae) from Japan, Scymnus ningshanensis Yu & Yao from China, and Harmonia axyridis (Pallas), a generalist species introduced from Asia that is currently widespread in eastern hemlock, Tsuga canadensis Carriere, forests. We measured the feeding preference of each beetle species when given the choice of A. tsugae and either 1) Pineus strobi (Hartig) on Pinus strobus L.; 2) Adelges laricis Vallot on Larix decidua Mill.; 3) Adelges cooleyi (Gillette) on Pseudotsuga menziesii (Mirb.) Franco; or 3) Paraprociphilus tessellatus (Fitch) on Alnus serrulata (Ait.) Willd. We evaluated beetle preference for adults, nymphs, and eggs of each prey species. Generally, when adult or nymphal prey stages were compared, S. tsugae preferred A. tsugae adults to P. strobi, A. cooleyi, A. laricis, and P. tessellatus. S. ningshanensis showed less preference between adelgid species, but it did not prefer P. tessellatus nymphs. When preferences for adelgid eggs were assayed, S. tsugae and S. ningshanensis showed no preference between A. tsugae and A. cooleyi or P. strobi, but S. tsugae did prefer A. tsugae to A. laricis. Larvae of S. tsugae were unable to survive on P. tessellatus nymphs. H. axyridis adults readily consumed both A. tsugae and P. tessellatus, but H. axyridis larvae did not complete their life cycle on A. tsugae. Our host range tests suggest that S. ningshanensis and S. tsugae may feed on several species of Adelgidae and that A. tsugae is often preferred.

Animals↗

Ion cycling in hemlock-northern hardwood forests of the southern Lake Superior region: a preliminary study.

Upland forests of the southern Lake Superior region are diverse and contain a shifting mosaic of eastern hemlock [Tsuga canadensis (L.) Carr.] and northern hardwood forests dominated by sugar maple (Acer saccharum Marsh.). In this study, we survey the relative effects of management practice (old growth vs. managed), forest cover type (hemlock vs. northern hardwood), and soil great group (Entic Haplorthod vs. Alfic Oxyaquic Fragiorthod) on ion cycling as a precursor to a longer-term, more detailed study. Bulk precipitation, throughfall, and soil leachates at three depths were collected for two growing seasons in eight stands on the Ottawa National Forest in the Upper Peninsula of Michigan. A total of 1210 solutions were analyzed for pH, Na, K, Mg, Ca, Cl, NO3, and SO4. Losses of base cations (Ca, Mg, K) and SO4 from the bottom of the rooting zone generally were greater in old-growth than in managed northern hardwoods on both fragic and nonfragic soils. Leaching losses of base cations and NO3 usually were greater beneath old-growth northern hardwoods than beneath old-growth hemlock on both soil types and for both forest cover types and management practices on fragic than nonfragic soils. Management practice, forest cover type, and soil type all appear to affect ion cycling within these forests. All of the stands featured striking losses of base cations that probably are influenced strongly by NO3 and SO4 in atmospheric deposition.

Acer↗

Radio ultrasound observations of the fetotoxic effects in sheep from ingestion of Conium maculatum (poison-hemlock).

Fetal movement in pregnant ewes gavaged with Conium maculatum (poison-hemlock) was reduced significantly, but temporarily. Fetal movement was observed by radio ultrasound at 45, 54 and 60 days of gestation in control ewes and on days 45, 54, and 60 of gestation immediately before and 1 hour following poison-hemlock feeding in treated ewes. Fetal movement was significantly reduced (P less than 0.01) 1 hour after poison-hemlock administration, but returned to normal within 18 hours post treatment. At parturition seven of eleven lambs born to seven treated ewes had varying degrees of front limb abnormalities. Modest to moderate flexure of the carpal joints, some lateral deviation in the front limbs at the pastern joint and kinked tails were observed. These malformations were transient and resolved spontaneously by 8 weeks after lambing.

Animals↗

Poison hemlock (Conium maculatum L.).

One of the most poisonous species amongst higher plants is Conium maculatum. It is a very common nitrophile weed species, belonging to the Apiaceae (formerly Umbelliferae) family. It contains some piperidine alkaloids (coniine, N-methyl-coniine, conhydrine, pseudoconhydrine, gamma-coniceine), which are formed by the cyclisation of an eight-carbon chain derived from four acetate units. gamma-Coniceine is the precursor of the other hemlock alkaloids. All vegetative organs, flowers and fruits contain alkaloids. The concentrations (both absolute and relative) of the different alkaloids depend on plant varieties, on ecological conditions and on the age of the plant. The characteristic biological effects of the plants are summarised on cattle, sheep, goat, swine, rabbit, elk, birds and insects and the symptoms of the human toxicosis (some cases of poisonings) are discussed according to the literature data. The general symptoms of hemlock poisoning are effects on nervous system (stimulation followed by paralysis of motor nerve endings and CNS stimulation and later depression), vomiting, trembling, problems in movement, slow and weak later rapid pulse, rapid respiration, salivation, urination, nausea, convulsions, coma and death.

Animals↗

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting.

Following planting, western hemlock (Tsuga heterophylla (Raf.) Sarg.) seedlings experience water stress and declining xylem pressure potential (Psi(x)). Low Psi(x) can result in xylem cavitation and embolism formation, causing a decline in hydraulic conductance. This study focused on the relationship between Psi(x), xylem cavitation and transpiration (E) of newly planted seedlings. Leaf specific hydraulic conductance (k(AB)) declined from 0.56 to 0.09 mmol m(-2) s(-1) MPa(-1) over a 9-day period. Stomatal conductance (g(s)) declined from 143.5 to 39.15 mmol m(-2) s(-1) over the same period without an associated change in environmental conditions. A vulnerability profile indicated a 30% loss in hydraulic conductivity when seedlings experienced a Psi(x) between -2.5 and -3.0 MPa. A Psi(x) of -4.0 MPa led to a complete loss of conductivity. We conclude that following planting, western hemlock seedlings often experience Psi(x) values that are low enough to cause xylem cavitation and a decline in k(AB).

Journal Article↗