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1.
The effect of intracellular calcium on the deformability of human erythrocytes was studied with a rheoscope, especially in relation to the dynamic structure of membrane cytoskeleton. The appropriate calcium-loading and calcium-depletion were performed to intact erythrocytes with A23187 in potassium buffer. The total calcium content was varied in the range of 0.25 to 3 times as much as normal content, without complete ATP depletion and shape change (the reduction of mean cell volume and the condensation of hemoglobin due to dehydration were avoided). Increasing the intracellular calcium content by about 1.5 times of normal, the deformability was distinctly decreased, while calcium depletion did not affect the deformability. Reduced deformability of the calcium-loaded erythrocytes was restored by the treatment with calmodulin inhibitors, W-7 or trifluoperazine. However, such an effect by calmodulin inhibitors was not detected on normal or calcium-depleted erythrocytes. In conclusion, the interaction between calcium-calmodulin complex and cytoskeletal proteins may affect the membrane stiffness which is regulated through the change of the cytoskeletal structure, and contributes to the deformability of erythrocytes.  相似文献   

2.
The deformability of human erythrocytes was measured in a rheoscope, as a function of intracellular calcium content (varied with ionophore (A23187) and CaCl2) without complete ATP depletion and echinocytic transformation. Loading calcium into intact erythrocytes (calcium content: 16.8 mumol/1 packed cells = 1.48 amol per cell), the cell volume and energy charge gradually decreased. Further, the membrane fluidity of the lipid portion decreased without crosslinking of membrane proteins. A distinct transition from deformable to undeformable cells was observed by the rheoscope technique: i.e., 50% transition occurred at 40-50 mumol calcium/1 packed cells (= 3.5-4.0 amol per cell) and more than 90% above 100 mumol/1 packed cells (= 6.5 amol per cell) at a shear stress of 140 dyn/cm2. The deformable cells maintained their deformability to ellipsoidal disks independent of the average calcium content. The underformable cells, separated as high-density cells by density gradient centrifugation after calcium-loading, showed lower glucose-6-phosphate dehydrogenase activity than low-density-deformable cells; thus, the calcium-loaded, undeformable cells were presumably in vivo aged cells. The younger cells, fractionated as low-density cells from intact erythrocytes, were more deformable than aged cells. Upon calcium-loading, the younger cells restored their cell volume and deformability, while the aged cells, containing originally more calcium and less ATP, decreased their volume and became undeformable. Therefore, calcium accumulation by ionophore-CaCl2 takes place in preference to aged cells of lower energy metabolism, and leads to cellular dehydration and loss of deformability, due to condensed hemoglobin and altered membrane organization.  相似文献   

3.
Maternal antibodies protect chicks from infection with pathogens early in life and may impact pathogen dynamics due to the alteration of the proportion of susceptible individuals in a population. We investigated the transfer of maternal antibodies against avian influenza virus (AIV) in a key AIV host species, the mallard (Anas platyrhynchos). Combining observations in both the field and in mallards kept in captivity, we connected maternal AIV antibody concentrations in eggs to (i) female body condition, (ii) female AIV antibody concentration, (iii) egg laying order, (iv) egg size and (v) embryo sex. We applied maternity analysis to the eggs collected in the field to account for intraspecific nest parasitism, which is reportedly high in Anseriformes, detecting parasitic eggs in one out of eight clutches. AIV antibody prevalence in free-living and captive females was respectively 48% and 56%, with 43% and 24% of the eggs receiving these antibodies maternally. In both field and captive study, maternal AIV antibody concentrations in egg yolk correlated positively with circulating AIV antibody concentrations in females. In the captive study, yolk AIV antibody concentrations correlated positively with egg laying order. Female body mass and egg size from the field and captive study, and embryos sex from the field study were not associated with maternal AIV antibody concentrations in eggs. Our study indicates that maternal AIV antibody transfer may potentially play an important role in shaping AIV infection dynamics in mallards.  相似文献   

4.
The deformability of human erythrocytes was investigated with a rheoscope to study the role of intracellular calcium in the dynamic cytoskeletal structure. Calcium was loaded to or depleted from erythrocytes with a calcium ionophore (A 23187) in a Na- or a K-HEPES buffer. (1) After calcium loading in the Na-HEPES buffer, the cell volume of erythrocytes was greatly reduced due to dehydration. On the contrary, upon calcium-loading or -depletion in the K-HEPES buffer, the intracellular calcium content could be varied in the range of 1/4 to 3 times as much as that of control cells without the reduction of mean cell volume. Further incubation without A 23187 and calcium in the K-HEPES buffer enabled the calcium-loaded erythrocytes to restore the cell shape and the ATP concentration. (2) When intracellular calcium content was increased to above 1.5 times of the normal value, the deformability was distinctly decreased. On the other hand, the deformability was unchanged when the intracellular calcium content was reduced below the normal level. (3) The deformability, once decreased due to the calcium accumulation, was recovered by the treatment with a calmodulin inhibitor, W-7 or trifluoperazine, while these drugs were not effective on the deformability of control or calcium-depleted erythrocytes. We conclude that the membrane stiffness which influence the deformability of erythrocytes, is modulated by the intracellular calcium content through the interaction between the calcium-calmodulin complex and the cytoskeletal proteins.  相似文献   

5.
S. JAMES REYNOLDS 《Ibis》2001,143(2):205-215
Many small passerines forage intensively for calcium-rich foods during laying. Increased incidences of shell defects in eggs of small passerines have been reported, particularly in western Europe, and these have been explained in terms of declining calcium availability in soils, resulting from prolonged anthropogenic acid deposition. Studies in the field have provided laying birds, nesting in areas of low calcium availability, with calcium supplements. An alternative approach was adopted in this study by allowing captive Zebra Finches Taeniopygia guttata to lay first clutches on ad libitum calcium, switching them to a low calcium diet for 72 hours for the formation of all but the first egg of the second clutch and reinstating ad libitum calcium for the final clutch. Control females had access to ad libitum calcium for all three clutches. Clutch sizes did not vary significantly between birds on low calcium and controls. The former took over three days longer to lay clutch 3 than did controls but the difference was not statistically significant. Birds on low calcium laid eggs that declined in shell ash mass with laying sequence, indicating that birds may have been calcium-limited. Although not statistically significant, eggshell thickness also declined with laying sequence in clutches laid by females on low calcium. The remaining egg measurements (shell mass, shell surface area and volume] of clutches laid by birds on low calcium did not differ significantly from those of controls. Furthermore, females on low calcium did not resort to skeletal reserves to provide sufficient calcium for egg formation. Dietary calcium appears to be of paramount importance in providing sufficient calcium for clutch formation.  相似文献   

6.
Pollution relative to phosphorus excretion in poultry manure as well as the soaring prices of phosphate, a non-renewable resource, remain of major importance. Thus, a good understanding of bird response regarding dietary phosphorus (P) is a prerequisite to optimise the utilisation of this essential element in broiler diets. A database built from 15 experiments with 203 treatments was used to predict the response of 21-day-old broilers to dietary non-phytate P (NPP), taking into account the main factors of variation, calcium (Ca) and microbial phytase derived from Aspergillus niger, in terms of average daily feed intake (ADFI), average daily gain (ADG), gain to feed (G:F) and tibia ash concentration. All criteria evolve linearly (P < 0.001) and quadratically (P < 0.001) with dietary NPP concentration. Dietary Ca affected the intercept and linear component for ADG (P < 0.01), G:F (P < 0.05) and tibia ash concentration (P < 0.001), whereas for ADFI, it affected only the intercept (P < 0.01). Microbial phytase addition impacted on the intercept, the linear and the quadratic coefficient for ADFI (P < 0.01), ADG (P < 0.001) and G:F (P < 0.05), and on the intercept and the linear component (P < 0.001) for tibia ash concentration. An evaluation of these models was then performed on a database built from 28 experiments and 255 treatments that were not used to perform the models. Results showed that ADFI, ADG and Tibia ash concentration were predicted fairly well (slope and intercept did not deviate from 0 to 1, respectively), whereas this was not the case for G:F. The increase in dietary Ca concentration aggravated P deficiency for all criteria while phytase addition had a positive effect. The more P deficiency was marked, the more the bird response to ADFI, ADG, G:F and tibia ash concentration was exacerbated. It must also be considered that even if the decrease in dietary Ca may improve P utilisation, it could in turn become limiting for bone mineralisation. In conclusion, this meta-analysis provides ways to reduce dietary P in broiler diets without impairing performance, taking into account dietary Ca and microbial phytase.  相似文献   

7.
Summary Calcium intake by wild rodents varies with season and habitat. This may have important ecological consequences; several studies have suggested that calcium availability may limit growth and reproduction. We studied the effect on growth of varying the calcium intake of captive wood mice Apodemus sylvaticus (L.) and bank voles Clethrionomys glareolus (Schreber 1780). To determine whether effects observed in the laboratory could be detected in free-living animals, we also compared the body weights and lengths of free-living animals on calcium-poor gritstone areas with those of rodents on calcium-rich limestone habitats where the dietary calcium concentration was between 2 and 5 times higher. Captive wood mice fed high calcium (0.87%) diet grew at the same rate as mice fed low calcium (0.30%) diet but continued growing for longer, thereby achieving higher asymptotic weights. In contrast, captive bank voles fed the high calcium diet grew more slowly and had lower asymptotic weights than voles fed the low calcium diet. As expected from the laboratory growth study, the higher calcium intake of free-living wood mice on the limestone was associated with greater body size compared with mice on the gritstone. However, bank voles were also larger on the limestone, even though high calcium intake impaired growth in captive animals. The contrast between wood mice and bank voles in the effects of calcium on growth, the reason why impaired growth may not be apparent in bank voles from calcium-rich habitats and the ecological significance of these results are discussed.  相似文献   

8.
Differences in growth conditions during early ontogeny have been suggested to cause permanent effects on the morphology and quality of birds. Yearly variation in growth conditions could thus result in morphological and quality differences between cohorts. In this study, we investigated the effect of small differences in the dietary protein content of captive ring-necked pheasants (Phasianus colchicus) during their first 8 wk posthatching. An experimental increase of the proportion of dietary protein during the first 3 wk of life accelerated growth, whereas a similar manipulation during the following 5 wk had only a limited effect. Compensatory growth during the postexperimental period equalized the size of chicks from different experimental treatments. However, a difference in tarsus length resulting from experimental treatment during the first 3 wk remained into adulthood. Furthermore, the protein content of the diet during the first 3 wk had an effect on the degree of fluctuating asymmetry in tarsus length, suggesting persistent effects on the quality of birds. The results of this study may explain size differences between cohorts that exist in pheasants and may also provide a link between the use of pesticides in agriculture and population effects on pheasants.  相似文献   

9.
Around 60% of pheasants released for shooting in the UK, an estimated 21 million birds, do not end up at their intended fate: being shot. This constitutes wastage, raising economic, environmental and ethical questions. We review what is known of the fates of released pheasants and consider why they do not directly contribute to the numbers harvested. We focus on four main explanations: predation, disease, starvation and dispersal, and highlight other important causes of mortality. For each explanation, we attempt to attribute levels of loss and identify timings or conditions when such losses may be heaviest. We review factors that exacerbate losses and methods available to mitigate them. Opportunities for amelioration may arise at all stages of the rearing and release of pheasants and involve changes to the conditions under which eggs are produced, the way young pheasants are reared or the management of the environment into which they are released. We found few studies investigating impacts of post-release management techniques on pheasant survival outside of the breeding season within a UK context. We found that a number of less commonly deployed practices focusing on early-life, pre-release management may improve survival. Given the scale of pheasant releasing in the UK, even improvements in survival of 1% would mean that ~?350,000 fewer birds die of natural causes. Complementing current post-release management with proven novel pre-release management interventions could reduce the number of pheasants required for release, whilst maintaining current shooting levels. Lowering release numbers would lower financial costs, benefit the environment and reduce some ethical concerns over the release and shooting of reared pheasants.  相似文献   

10.
Transfer coefficients (the equilibrium ratio be-tween radionuclide activity concentration in milk or meat and the daily intake of radionuclide) are widely used to predict the contamination of animal products following the release of radionuclides into the environment. For a transfer coefficient to be generally applicable, its value must be constant for a range of circumstances. However, this is not the case for radiostrontium, the behaviour of which is strongly influenced by that of the homeostatically controlled nutrient, calcium. In this study, a relationship is derived between radiostrontium transfer coefficients and dietary calcium intake which takes into account the observed ratio for strontium:calcium transfer to milk. This relationship is tested against a range of observed data collated from the literature (n = 30) and found to account for 93% of the variability in transfer coefficient values. Model calculations show that a reduction in F m of at least 40%–60% would be expected if dairy cattle, fed rations typical for well-managed herds, were supplemented with 100–200 g per day. Larger reductions would be predicted when dietary calcium intake is low. Received: 14 October 1996 / Accepted in revised form: 26 November 1996  相似文献   

11.
Fat sand rats Psammomys obesus feed exclusively on plants of the family Chenopodiaceae, which contain high concentrations of chloride salts (NaCl, KCl) and oxalate salts. Ingestion of large quantities of oxalate is challenging for mammals because oxalate chelates Ca(2+) cations, reducing Ca(2+) availability. Oxalate is a metabolic end-point in mammalian metabolism, however it can be broken-down by intestinal bacteria. We predicted that in fat sand rats microbial breakdown of oxalate will be substantial due to the high dietary load. In addition, since a high concentration of soluble chloride salts increases the solubility of calcium oxalate in solution, we examined whether a change in the intake of chloride salts affects microbial oxalate breakdown and calcium excretion in fat sand rats. We measured oxalate, calcium and other inorganic matter (ash) intake and excretion in fat sand rats feeding on two different diets: saltbush (Atriplex halimus), their natural diet, and goose-foot (Chenopodium album), a non-native chenopod on which fat sand rats will readily feed and that has a similar oxalate content to saltbush but only 2/3 of the ash content. In animals feeding on both diets, 65-80% of the oxalate ingested did not appear in urine or faeces. In animals consuming the more saline saltbush, significantly more oxalate was apparently degraded (p<0.001), while significantly less oxalate was excreted in urine (p<0.01) and in faeces (p<0.05). We propose, therefore, that fat sand rats rely on symbiotic bacteria to remove a large portion of the oxalates ingested with their diet, and that the high dietary salt intake may play a beneficial role in their oxalate and calcium metabolism.  相似文献   

12.
Phytase has long been used to decrease the inorganic phosphorus (Pi) input in poultry diet. The current study was conducted to investigate the effects of Pi supplementation on laying performance, egg quality and phosphate–calcium metabolism in Hy-Line Brown laying hens fed phytase. Layers (n = 504, 29 weeks old) were randomly assigned to seven treatments with six replicates of 12 birds. The corn–soybean meal-based diet contained 0.12% non-phytate phosphorus (nPP), 3.8% calcium, 2415 IU/kg vitamin D3 and 2000 FTU/kg phytase. Inorganic phosphorus (in the form of mono-dicalcium phosphate) was added into the basal diet to construct seven experimental diets; the final dietary nPP levels were 0.12%, 0.17%, 0.22%, 0.27%, 0.32%, 0.37% and 0.42%. The feeding trial lasted 12 weeks (hens from 29 to 40 weeks of age). Laying performance (housed laying rate, egg weight, egg mass, daily feed intake and feed conversion ratio) was weekly calculated. Egg quality (egg shape index, shell strength, shell thickness, albumen height, yolk colour and Haugh units), serum parameters (calcium, phosphorus, parathyroid hormone, calcitonin and 1,25-dihydroxyvitamin D), tibia quality (breaking strength, and calcium, phosphorus and ash contents), intestinal gene expression (type IIb sodium-dependent phosphate cotransporter, NaPi-IIb) and phosphorus excretion were determined at the end of the trial. No differences were observed on laying performance, egg quality, serum parameters and tibia quality. Hens fed 0.17% nPP had increased (P < 0.01) duodenum NaPi-IIb expression compared to all other treatments. Phosphorus excretion linearly increased with an increase in dietary nPP (phosphorus excretion = 1.7916 × nPP + 0.2157; R2 = 0.9609, P = 0.001). In conclusion, corn–soybean meal-based diets containing 0.12% nPP, 3.8% calcium, 2415 IU/kg vitamin D3 and 2000 FTU/kg phytase would meet the requirements for egg production in Hy-Line Brown laying hens (29 to 40 weeks of age).  相似文献   

13.
The phenomenon of dart shooting in several species of land snailshas still not been explained. We were interested in whetherthe dart can function as a nuptial gift of calcium, as previouslyproposed. Donating calcium would increase the fitness of theoffspring and thereby result in a higher reproductive successfor the donor. We confirmed in Helix aspersa that the developingembryo takes up calcium from the egg shell for the formationof its embryonic shell. However, other results from behaviouralobservations and calcium measurements in various reproductivestructures do not support the calcium hypothesis. We found thatthe dart penetrated the skin in 91.7% or the shootings, butit was internalized by the recipient in only 6.3% of the shootings.The amount of calcium in one dart is roughly equal to that ofone egg, and thus it would not contribute significantly to anaverage clutch of 59 eggs. The spermatophore contains virtuallyno calcium, and therefore it is unlikely that the dart signalsa donation of calcium with the sperm. The dart is also unlikelyto influence egg laying since dart shooting does not predicteither the latency or the productivity of egg laying in theshooter or the recipient. We conclude that the love dart ofHelix aspersa is not a gift of calcium. Instead, we suggestthat it is a vehicle to introduce a substance into the partnerto influence the fate of the donated sperm. (Received 8 November 1996; accepted 25 April 1997)  相似文献   

14.
模拟黄腹角雉野外生存环境和自然种群结构,采用较大空间散放与多个体群养相结合的方法进行饲养试验,其笼养繁殖性能比对照组传统对养方式显著提高.通过两年来的试验研究,结果表明:试验组平均产卵量为8.5枚,卵受精率为82.6%,受精卵孵化率为88.7%;对照组平均产卵量5.7枚,卵受精率50.3%,受精卵孵化率72.4%,试验组明显高于对照组.同时对两组间的卵重、卵形、产卵期进行了测定,无显著差异。  相似文献   

15.
Changes in dietary preferences in animal species play a pivotal role in niche specialization. Here, we investigate how divergence of foraging behaviour affects the trophic position of animals and thereby their role for ecosystem processes. As a model, we used two closely related bat species, Myotis myotis and M. blythii oxygnathus, that are morphologically very similar and share the same roosts, but show clear behavioural divergence in habitat selection and foraging. Based on previous dietary studies on synanthropic populations in Central Europe, we hypothesised that M. myotis would mainly prey on predatory arthropods (i.e., secondary consumers) while M. blythii oxygnathus would eat herbivorous insects (i.e., primary consumers). We thus expected that the sibling bats would be at different trophic levels. We first conducted a validation experiment with captive bats in the laboratory and measured isotopic discrimination, i.e., the stepwise enrichment of heavy in relation to light isotopes between consumer and diet, in insectivorous bats for the first time. We then tested our trophic level hypothesis in the field at an ancient site of natural coexistence for the two species (Bulgaria, south-eastern Europe) using stable isotope analyses. As predicted, secondary consumer arthropods (carabid beetles; Coleoptera) were more enriched in 15N than primary consumer arthropods (tettigoniids; Orthoptera), and accordingly wing tissue of M. myotis was more enriched in 15N than tissue of M. blythii oxygnathus. According to a Bayesian mixing model, M. blythii oxygnathus indeed fed almost exclusively on primary consumers (98%), while M. myotis ate a mix of secondary (50%), but also, and to a considerable extent, primary consumers (50%). Our study highlights that morphologically almost identical, sympatric sibling species may forage at divergent trophic levels, and, thus may have different effects on ecosystem processes.  相似文献   

16.
Energy maximization, time minimization, and linear programming models subject to various constraints have dominated foraging ecology ideas and methods for decades. However, animals must use very complex physiological processes and foraging decisions to ensure fitness that in many cases may not be adequately described by these approaches. An example of this problem occurs when brown bears, Ursus arctos, have access to both abundant salmon and fruit. Salmon are one of the most energy and nutrient dense foods available to bears. Fruits are often high in soluble carbohydrates, low to deficient in many required nutrients, and more difficult to efficiently exploit. Therefore, wild brown bears that fatten primarily on fruits without access to salmon are 50% smaller than salmon-feeding bears. Thus, we predicted based on a linear, energy-maximizing model without dietary interaction effects that wild brown bears with access to both abundant salmon and fruit would feed almost exclusively on salmon. However, wild adult females with or without accompanying offspring foraged three times longer per day on fruit than on salmon. Similarly, the relative dry matter intake of ad libitum apples and salmon by captive, adult brown bears averaged 76±5% fruit and 24±5% salmon. Captive brown bears consuming mixed diets with intermediate dietary protein levels had 60% lower maintenance energy costs, 37% to 139% higher efficiencies of mass gain, and 72% to 520% higher maximum rates of gain than when they consumed either salmon or fruit alone. These relationships were nonlinear functions of dietary protein content in which salmon and fruit provided complementary nutritional resources. Both wild and captive bears attempted to regulate total protein, energy, and carbohydrate intake within a multidimensional intake target that both maximized energy intake and mass gain.  相似文献   

17.
The regulatory protein, calmodulin, undergoes major conformational changes in response to changes in intracellular calcium concentration. Furthermore, calmodulin has been reported to have lysine residues which markedly increase their reactivity toward electrophilic substances in the calcium-loaded state. We found that calmodulin formed two to three times more stable adducts with acetaldehyde in the calcium-loaded state as compared to the calcium-free state. Competition-binding studies showed that calmodulin could preferentially compete with albumin for acetaldehyde in the presence, but not in the absence, of calcium. When calmodulin was in the calcium-loaded state, trifluoperazine, an inhibitor of calmodulin activity, significantly decreased the stable binding of acetaldehyde to the protein, whereas in the calcium-free state, minimal effects on binding were observed. Since calmodulin is involved in regulation of multiple important processes in the cell, it is possible that acetaldehyde-calmodulin adducts could contribute to liver injury by perturbation of calcium-dependent homeostatic mechanisms within the hepatocyte.  相似文献   

18.
Social tolerance crucially affects the life of group‐living animals as it can influence, among other things, their competitive regimes, access to food, learning behavior, and recruitment. However, social tolerance tests were mainly conducted in semi‐free or captive populations, and we know little about the behavioral mechanisms and consequences of social tolerance under natural conditions. We therefore developed a co‐feeding experiment to measure social tolerance in groups of wild and captive animals across two primate species. Specifically, we recorded the social tolerance level of redfronted lemurs (Eulemur rufifrons, four wild, one captive group) and ringtailed lemurs (Lemur catta, three wild, three captive groups) by presenting a clumped food resource in an experimental arena, and recorded patterns of resource use during the experiment. Because redfronted lemurs exhibit lower levels of decided conflicts than ringtailed lemurs, we predicted that they would be socially more tolerant. The probability for an individual to feed in the arena was higher in redfronted lemurs than in ringtailed lemurs. In addition, in both species, the probability for an individual to feed in the arena was higher in the captive populations than in their wild counterparts, suggesting that proximate factors, such as a relaxation of feeding competition in captivity, may adapt species‐specific levels of social tolerance to local levels of food availability. Hence, the number of individuals co‐feeding on a valuable food resource appears to be a useful proxy of social tolerance that could be measured with this experimental setup in other wild and captive species as well.  相似文献   

19.
Based on Trivers' (1974) theory of parent/offspring conflict, we predicted that weaning would be accelerated under poor dietary conditions in an aseasonally breeding, monomorphic ungulate, the collared peccary (Tayassu tajacu). Suckling behavior was observed in 18 captive litters under a dietary treatment of high or low protein formulated to mimic forage under ecological conditions of normal rainfall vs. drought. Litters on the low protein diet weaned later (mean of 27 weeks) than litters on the high protein diet (mean of 21 weeks). Other indices of cumulative suckling effort (weaning weight, cumulative minutes suckling, cumulative bouts) did not differ between dietary treatments. No differences in suckling effort were found relative to offspring sex or litter size. Milk did not completely compensate for the low protein diet affecting growth of young in the second and third phases of lactation. Some nursers did not conserve bodily resources as would be expected if the benefits of short interbirth intervals outweighed the costs of lactation during unpredictable drought conditions.  相似文献   

20.
Summary The nutrient content and balance of the horsetail Equisetum fluviatile was measured in samples taken from southern James Bay during early spring, summer and fall, and related to the nutrient requirements of geese at these times. Young emergent stems contained in excess of 20% protein of well balanced amino acid composition, apart from deficiencies of cystine and methionine. The ash fraction contained high levels of calcium, magnesium, potassium and phosphorus, which would satisfy the mineral needs of breeding and growing geese. The aerial parts of E. fluviatile did not contain much energy as neutral fats and soluble carbohydrate. The growth pattern of E. fluviatile, with young aerial stems arising throughout the summer, would provide a source of succulent, high-protein food. This could be used by geese migrating to the breeding grounds, during egg formation, to facilitate growth of goslings, and in the reacquisition of body protein lost during reproduction. The nutritive value of E. fluviatile was not high during the fall.  相似文献   

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