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1.
Electron microscopical observations on algal samples collected in 1992 in the middle or final stages of a red tide in Hiroshima Bay, Japan, revealed virus-like particles (VLPs) in the red tide alga Heterosigma akashiwo (Class Raphidophyceae). The host cells appeared moribund and the VLPs were located in and around the nuclear area. The VLPs were icosahedral, ca. 185 nm in diameter, and generated from the periphery of several viroplasms. VLPs were also observed in three other types of H. akashiwo-like cells, which were morphologically distinguishable from each other. The appearance of VLPs in the red tide alga could explain the dramatically rapid termination of this red tide.  相似文献   

2.
I. Imai 《Marine Biology》1989,103(2):235-239
Cyst formation of the noxious red tide flagellate Chattonella marina (Raphidophyceae), cultured from a cyst in sediments of Suo-nada, Japan in 1986, was investigated by culture experiments. Nitrogen limitation was effective in inducing cyst formation. After incubation in N-limited medium at 25°C with 10000 lux on a 14 h light, 10 h dark photo-cycle, cultures were subjected to six different light intensities (0 10000 lux) in order to determine effects of light intensity on cyst formation. Cysts were formed under light intensities of 1000 lux or below, but not at 10000 lux. Cysts formed in culture displayed morphological characteristics quite similar to those natural cysts observed in sediments collected from the Inland Sea of Japan. Observations on nucleus, using diaminidinophenylindole (DAPI)-staining and epifluorescence microscopy, indicated that the majority of cysts formed in culture were uninucleate, a few were binucleate. Germination of cysts was observed under adequate conditions for germination (22°C, 3500 lux with a 14 h light, 10 h dark photo-cycle) even after storage at 11°C in the dark for more than 4 mo.  相似文献   

3.
S. Yamochi  T. Abe 《Marine Biology》1984,83(3):255-261
The ecological role of diel vertical migration of Heterosigma akashiwo Hada to initiate red tide was investigated in Tanigawa Fishing Port and Sano Harbor, Osaka Bay, Japan during red tide seasons in 1979 and 1980. This species migrated toward the surface early in the morning at a velocity of 1.0 to 1.3 m h-1. Downward migration was found in the afternoon, and more than 2.0×103 cells ml-1 aggregated in the bottom layer at night. The upward migration started before sunrise and downward shifting occurred prior to sunset. The movement is presumably associated with circadian rhythm, which is known as one of the biological periodicities. H. akashiwo crossed steep temperature and salinity gradients (6.5°C and 5.7 S) during the diel vertical migration. High values of particulate organic carbon and nitrogen concentrations were obtained in dialysis bags suspended in situ at identical layers with high cell concentration, while the values for surface and bottom bags were comparatively low. The results reveal that H. akashiwo migrates toward the sea surface to carry out photosynthesis effectively, and to the bottom to utilize nutrients efficiently.  相似文献   

4.
L. B. Connell 《Marine Biology》2000,136(6):953-960
The internal transcribed spacer (ITS) region from 19 isolates of the algal genus Heterosigma (Chromophyta: Raphidophyceae) was amplified by polymerase chain-reaction (PCR) and sequenced. Isolates were obtained from both the Atlantic and Pacific basins, including Europe, eastern North America, western North America, Japan and New Zealand. This study presents evidence that all Heterosigma isolates in this study are representatives of one species (H. akashiwo). All 19 isolates, except one (LB 2005) had identical ITS sequence (98.31% similar by pairwise comparison); Isolate LB 2005 may represent a separate subspecies. Such high degree of ITS sequence identity implies that the organism has spread between oceanic regions in geologically recent times, possibly by human means. In addition to those from Heterosigma spp., the ITS regions from other marine Raphidophyceae (Chattonella antiqua, C. marina, C. subsalsa, Fibrocapsa japonica, and Olisthodiscus luteus) were amplified and sequenced using PCR. Total ITS lengths differed among the Raphidophyceae (C. antiqua, 577 base pairs (bp); C. marina, 577 bp;. C. subsalsa, 579 bp; F. japonica, 830 bp; H. akashiwo, 561 and 563 bp; O. luteus, 829 bp), but 5.8S rDNA sequences were similar in size (13 to 142 bp). The high ITS sequence identity between C. antiqua and C. marina (>99.9% by pairwise comparison) suggests the need for a taxonomic review of these species encompassing all morphological, genetic, physiological and biochemical information. Additionally, a number of cultures of Raphidophyceae were positively identified. In general, ITS comparisons among the Raphidophyceae may be most useful at the level of species determination rather than at the population level. Received: 12 July 1999 / Accepted: 16 March 2000  相似文献   

5.
Toxic mechanisms of the red tide flagellate, Chattonella marina, collected in 1985 from Kagoshima Bay, Japan, were studied at the subcellular level. C. marina was found to reduce ferricytochrome c at a rate related to the concentration of plankton cells. Ca. 50% of the cytochrome c reduction was inhibited by the addition of 100 U superoxide dismutase ml-1. These results suggest that a part of the cytochrome c reduction was caused by a superoxide anion which was extracellulary released from C. marina. Moreover, a small amount of hydrogen peroxide was detected in the C. marina suspension using the fluorescence spectrophotometric assay method. The identity of the hydrogen peroxide was confirmed by its reaction with 500 U catalase ml-1. It is thus proposed that C. marina produces harmful active oxygen radicals and therefore exhibits a toxic effect on surrounding living organisms. In agreement with these results, C. marina strongly inhibited the proliferation of marine bacteria, Vibrio alginolyticus, in a plankton/bacteria co-culture system. The growth inhibition of bacteria caused by C. marina was related to the density and the metabolic potential of C. marina. Ruptured plankton showed no toxic effect on the bacteria. Furthermore, the toxic effect of C. marina on V. alginolyticus was completely suppressed by the addition of catalase and superoxide dismutase. In addition to these radical-scavenging enzymes, a chemical scavenger, sodium benzoate, also had a protective effect. These results suggest that oxygen radicals are important in the toxic action of C. marina.  相似文献   

6.
A marine algicidal gliding bacterium Cytophaga sp. strain J18/M01 was isolated in 1990 from a station in northern Harima-Nada, the Seto Inland Sea, Japan, using the harmful red tide alga Chattonella antiqua (Hada) Ono as a susceptible organism. The bacterium can prey upon various species of microalgae. Temporal fluctuations of this bacterium and Chattonella spp. [C. antiqua and C. marina (Subrahmanyan) Hara et Chihara] were investigated weekly at the above station in the summer of 1997 and 1998, using immunofluorescence assay employing highly specific polyclonal antibodies for the bacterium. In the summer of 1997, the cell density of Chattonella spp. showed a maximum value (70 cells ml−1) on 8 July, and decreased thereafter. The bacterium Cytophaga sp. J18/M01 was commonly detected around a few hundreds of cells per milliliter or less. The number of Cytophaga sp. J18/M01 increased after the peak of Chattonella spp., and the maximum cell number of the bacterium was 1350 ml−1. This algicidal bacterium also followed the changes of total amounts of microalgal biomass (chlorophyll a+pheophytin) when Chattonella spp. were absent. In the summer of 1998, Chattonella spp. were relatively less abundant (maximum 21 cells ml−1), and the algicidal bacterium Cytophaga sp. J18/M01 showed a close relationship with the change of total microalgal biomass. The present study suggests that the algicidal bacterium Cytophaga sp. J18/M01 preyed upon, not only harmful red tide microalgae, but also other common microalgae such as diatoms, and the bacterium presumably plays an important role in regulating microalgal biomass in natural marine environments. Received: 20 April 2000 / Accepted: 1 December 2000  相似文献   

7.
In an attempt to clarify the surface structure of Heterosigma akashiwo, the organisms were processed by a series of currently utilized methods for carbohydrate and protein histochemistry of light microscopy. The results indicated that: (1) H. akashiwo cell has previously undetected glycocalyx on the cell surface; (2) this extracellular structure consists of at least two different types of acidic complex carbohydrates, sulfated and non-sulfated, together with a neutral carbohydrate-protein complex with 1,2-glycol groups and -d-mannosyl and -d-glucosyl residues; (3) a particular moiety of acidic complex carbohydrates involved in the glycocalyx is presumably hyaluronic acid or a closely related substance; and (4) the histochemical nature of the glycocalyx on the cell surface of the present species is comparable with that of Chattonella antiqua, excluding a difference in the nature of the neutral carbohydrate-protein complex.  相似文献   

8.
Although the ichthyotoxic mechanism of Chattonella marina is still unknown, several lines of evidence suggest that the reactive oxygen species (ROS), such as superoxide anion (O2-), hydrogen peroxide (H2O2), and hydroxyl radical (·OH), produced by C. marina are involved in the mortality of fish exposed to this flagellate. Recently, we found that the cell-free supernatant prepared from C. marina, which is considered to contain the glycocalyx, showed NADPH-dependent O2- generation. In this study, we prepared antiserum against the crude glycocalyx of C. marina. Using indirect immunofluorescence, it was confirmed that the antiserum specifically reacted with C. marina cells. In addition to C. marina, the antiserum also reacted with other raphidophycean flagellates such as Heterosigma akashiwo, Olisthodiscus luteus, and Fibrocapsa japonica, whereas no reactivity was observed against six other flagellate species tested. These results suggest that raphidophycean flagellates have common epitopes recognized by the antiserum. Interestingly, immunohistochemical analysis of paraformaldehyde-fixed gill lamellae from yellowtail exposed to C. marina revealed that the antiserum stained the surface of gill lamellae, while no such staining pattern was observed in control gill lamellae. These results suggest that the glycocalyx may be discharged when C. marina cells are inhaled into the fishes' mouths and then come into contact with the gill surface. Based on the present results, together with our previous findings, we propose that continuous accumulation of the discharged glycocalyx on the gill surface occurs during C. marina exposure, which may be responsible for the ROS-mediated severe gill tissue damage leading to fish death.  相似文献   

9.
Life cycle changes that allow populations of the toxic dinoflagellate Gonyaulax tamarensis Lebour to inhabit the benthos and the plankton alternately are important factors regulationg the initiation and decline of blooms in restricted embavments. When the dynamics of these estuarine populations were monitored during “bloom” and “non bloom” years, it was shown that: (1) each year, germination of benthie cysts inoculated the overlying waters during the vernal warming period, but a large residual population remained in the sediments throughout the blooms; (2) the resulting planktonic population began growth under suboptimal temperature conditions; (3) the populations developed from this inoculum through asexual reproduction until sexuality (and cyst formation) were induced; (4) encystment was not linked to any obvious environmental cue and occurred under apparently optimal conditions; and (5) an increase in the number of non-mitotic swimming cells (planozygotes, the precursors to dormant cysts) accompanied the rapid decline of the planktonic population. Thus encystment, in combination with hypothesized losses due to advection and grazing, contributed substantiatly to the decline of the vegetative cell population. We conclude that the encystment/excystment cycle temporally restricts the occurrence of the vegetative population and may not be optimized for rapid or sustained vegetative growth and bloom formation in shallow embayments. The factors that distinguish “bloom” from “non-bloom” years thus appear to be operating on the growth of the planktonic population.  相似文献   

10.
Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey (Alouatta seniculus) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km2, increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.  相似文献   

11.
The seasonal behavior of both vegetative cells and cysts of dinophytes Scrippsiella spp., mostly S. trochoidea, which is the dominant group among dinoflagellate populations in Onagawa Bay on the northeastern coast of Honshu, Japan, was investigated between 1990 and 1992. The germination of the cysts after 8 d incubation under favorable laboratory conditions was examined using the extinction dilution method. Incessant germination occurred throughout the year, but the germination ratio (no. of germinable cysts/total cysts) varied seasonally with a marked fluctuation during summer when vegetative cells in the water column were abundant. Although such fluctuation largely reflects the variable flux of newly deposited immature cysts produced by the vegetative cells, the regulation of germination caused by a lowered saturation of dissolved oxygen (DO) under thermally stratified conditions was also suggested. During winter, while the cysts germinated in the laboratory, vegetative cells were not found in the water column. These facts suggest that germination in situ is regulated by low temperature in winter and possibly by lowered DO and by cyst age as well in summer. Such regulation prevents simultaneous germination of all the cysts, which is disadvantageous for the population because it would be more difficult to survive adverse conditions such as successive nutrient depletion and higher grazing risk.  相似文献   

12.
The structure and some functional characteristics of the plankton community at the time of aExuviaella cordata red tide were investigated in Burgas Bay, Bulgaria, in May and June 1986. Characteristics of main plankton components (phyto-, bacterio-, nanoheterotrophic plankton, ciliates and mesoplankton) in the bloom area are presented. Development of theE. cordata bloom was determined by abiotic conditions among which eutrophication and salinity decrease caused the patchy character of its rapid development. Attainment of maximum red tide (ca. 1x109 cells l-1; 1x103 g m-3) from background (500 to 800x103 cells l-1; 600 mg m-3) took 3 to 7d. Growth rate () during that period was 1.2 to 2.2 doublings per day. A sharp bloom decline (3 to 4d) was caused by parasitic flagellates destroying the alga's chloroplast. Diel biomass losses due to grazing remained below 5%. Metabolites and degradation products ofE. cordata revealed no pronounced toxic effects on the other components of the planktonic community. The rapid bloom degradation due to the effects of parasitic flagellates indicates the high potential of ecosystem self-regulation.  相似文献   

13.
Brachiomonas submarina Bohlin (Chlorophyceae), a euryhaline marine flagellate, can osmoregulate over a wide range of external salinity. The alga exhibits maximum water content at 100% artificial seawater (ASW), and shows only a small water loss (<15%) when salinity is increased to 300% ASW. The non-aqueous volume of the cells is increased at salinities higher than 100% ASW. This is partially attributable to the accumulation of glycerol. Glycerol is the major osmoregulatory organic solute in this flagellate. The alga also shows an accumulation of amino acids in response to increased salinity. The contribution of glycerol and amino acids to intracellular osmolarity is only 9% at 10% ASW, but accounts for 49% at 300% ASW. The remainder of the osmotic balance is due to uptake and accumulation of inorganic ions, particularly sodium, potassium and chloride.  相似文献   

14.
This study documents the effects of two consecutive disturbances on coral community structures in the Gulf of Oman (United Arab Emirates); Cyclone Gonu in June 2007 and the Cochlodinium polykrikoides harmful algal bloom (HAB) that persisted from August 2008 until May 2009. Coral cover, colony densities, size class frequency distributions, and geometric growth rates derived from size class transition probability matrices were used to assess the post-Gonu and post-HAB recovery trajectories at four sites. The net effects of these disturbances were fourfold: (i) storm damage caused >50% losses of live branching and tabular coral cover by fragmentation and dislodgment of pocilloporid and acroporid colonies; (ii) Pocillopora damicornis colonies that survived the cyclone experienced mass mortality during the first 3?months of the HAB, resulting in localized extirpation of this species; (iii) variable Acropora mortality during the HAB indicated individual colony, rather than taxa-wide, susceptibility; and (iv) massive colony coral taxa were resistant to both disturbances.  相似文献   

15.
Iron-stress-mediated effects on biochemical constituents of the red tide dinoflagellateGymnodinium sanguineum Hirasaka were examined in 1988 by comparing Fe-replete and Fe-deplete batch cultures. The influence of nitrogen source (NO3 or NH4) on characteristics of Fe-deplete cells was also studied [i.e., Fe-deplete/NO3-grown (= — Fe/NO3) vs Fe-deplete/NH4-grown (= — Fe/NH4)]. Common to both N sources were reductions of chlorophylla (chla) and Fe quotas (per cell volume) by 75% and ca. 1.5 orders of magnitude, respectively, under Fe depletion. The Fe requirement ofG. sanguineum exceeded those of certain neritic diatoms by one to two orders of magnitude. — Fe/NH4 cells exhibited 30 to 50% greater N quotas and free amino acid:protein ratios than did Fe-deplete cells grown on NO3. In vivo fluorescence:chla increased with Fe deficiency particularly in — Fe/NO3 cultures, surpassing — Fe/NH4 values by ca. two-fold. Effects of Fe depletion were consistent with this element's essential role in the biosynthesis of chla and components of the photosynthetic electron transport (PET) system, and also in NO3 utilization. Fe:N ratios were larger (1.5-fold) for iron-deficient NO3-grown than NH4-grown cells, likely reflecting the Fe content of NO3 assimilatory enzymes [nitrate (NR) and nitrite (NiR) reductase] and of electron transport components needed to provide reductant, coupled with a diminished capacity of — Fe/NO3 cells to acquire and assimilate nitrogen. Indicators of PET efficiency suggested that under iron stress, supply of Fe for NR and NiR is partly at the expense of iron-containing PET components. Utilization of nitrate by NO3-grown cells was inhibited sufficiently by Fe depletion to yield symptoms bordering on N deficiency. In an ecological context, the most important effect mediated by nitrogen source may be the determination of critical QFe (i.e., Fe required to just sustain maximal growth), thereby regulating the degree of growth limitation for a given subsaturating iron concentration.  相似文献   

16.
G. W. Lopez 《Marine Biology》1982,68(3):333-341
Population dynamics of a benthic harpacticoid copepod (Tisbe cucumariae) were studied in a seawater-system holding tank during fall-spring, 1979/1980, and January, 1981. These populations were opoortunistic scavengers, especially on the dead bodies of Ciona intestinalis, a solitary tunicate. Copepod population densities, life-history stage structures, sex ratio and percentages of females with egg sacs were determined in the absence or in the presence of dead tunicates (from death through dissolution to final disappearance). Taken together, all the data are consistent with a hypothesis for food-dependent cycling of populations of T. cucumariae through the following 4 density states: (1) a maintenance state of low density, which is the usual condition during the absence of a concentrated food source; (2) a colonization state of moderate density, which results from attraction and immigration to a new food source; (3) a bloom state at the food source, where a high density results from intensive reproduction combined with a tendency of all life stages to remain at the site; (4) a dispersal state of moderate density as the copepods emigrate from the site of the consumed food. Evidence is also presented for a facultative slowing of naupliar development in T. cucumariae in the absence of a concentrated food source such as a dead tunicate. Possible advantages afforded by delayed development are discussed in terms of risk-spreading in a stochastically hostile environment.  相似文献   

17.
海洋赤潮藻球形棕囊藻在氮磷富营养下的细胞增殖   总被引:1,自引:0,他引:1  
利用常见海洋赤潮微藻球形棕囊藻(Phaeocystis globosa)为试验研究材料,以舵海洋微藻营养液为对照(1P1N:磷质量浓度为5×10-3g·L-1.氮质量浓度为75×10-3g·L-1),设置3组富磷和富氮营养处理(3P1N:磷质量浓度为15×10-3 g·L-1,氮质量浓度为75×10-3 g·L-1;1P3N:磷质量浓度为5××10-3 g·L-1,氮质量浓度为225×10-3 g·L-1;3P3N:磷质量浓度为15××10-3 g·L-1,氮质量浓度为225×10-3 g·L-1),利用细胞记数和叶绿素荧光测定等方法研究了藻细胞在不同富磷和富氮条件的增殖情况.结果显示,不同浓度磷和氮营养下的藻体荧光值变化在试验周期内均呈现"S"型曲线,表明藻细胞的生长经历缓慢期,快速期和平缓期3个阶段;同时,不同的富磷和富氮营养条件对球形棕囊藻的叶绿素荧光值有一定的影响,其中在对照1P1N下的藻体荧光值最低,在试验结束时(第10天)只有850 μg·L-1,而在3P1N,1P3N和3P3N条件下的藻体荧光值均达到900 μg·L-1以上,显著高于1P1N下的藻体荧光值,表明富磷和富氮营养可以促进藻细胞的生长增殖,但在试验设置的不同富磷和富氮营养下的藻体荧光值之间没有显著的差异.就不同磷和氮营养条件下的藻最大比生长速率而言,3P3N和3P1N条件下的最大,均达到0.77 d-1,明显高于1P1N和1P3N条件下的藻最大比生长速率(分别只有0.70 d-1和0.69 d-1).此外,试验结束时细胞密度的变化趋势与藻体荧光值相似,富磷和富氮营养条件下的细胞密度显著高于1P1N下的细胞密度,而富磷和富氮营养条件下的细胞密度间也不存在显著的差异.研究结果揭示,水体中的高磷和高氮营养浓度是导致藻细胞大量快速增殖的一个主要因素,而利用叶绿素荧光来测定藻细胞增殖是一种快速、简便,灵敏和可靠的方法,可在今后赤潮监测过程中多加利用,以能及时、准确地预测预报赤潮爆发.从而减少其对环境和经济的影响.  相似文献   

18.
P. O. Yund  A. Stires 《Marine Biology》2002,141(5):955-963
Recent interest in the dynamics of marine invertebrate populations has focused largely on taxa with an open population structure. However, in many colonial taxa with limited larval dispersal, settlers may be locally derived. Consequently, dynamics may vary among sites that are separated by relatively short distances. This study explored spatial variation in temporal dynamics of colonial ascidians (Botryllus schlosseri Pallas) inhabiting five sites distributed along a ≈ 17-km temperature and phytoplankton gradient in the Damariscotta River estuary, Maine, USA. Settlement and population densities and sexual reproductive status were assayed throughout the summer seasons of 1996 and 1997. Sexual reproduction and larval settlement commenced earlier in the summer in up-river populations, which subsequently underwent a seasonal population explosion that was much smalier in down-river populations. Two peaks in settlement density up-river (in early July and early September) suggest that colonies there may have completed two sexual generations, in contrast to a single generation at down-river sites. Similar spatial variation is expected among populations of other taxa with limited larval dispersal when they are distributed across environmental gradients. Published online: 18 September 2002  相似文献   

19.
Picocyanobacteria, which specifically excited the phycobilin pigments by green light, were numerically counted under an epifluorescence microscope in the Kuroshio and the Oyashio between 1 and 9 July 1983, and in coastal waters between 10 and 12 August 1983 off Japan. The fluorescence, and the various morphological and chemical characters of the picocyanobacteria were evaluated by using monospecific strains isolated from the study area. Plasma volume determined on the epifluorescence microscopic photographs was converted into cellular organic carbon using a relation obtained from the isolates. Percentages of the picocyanobacteria biomass in terms of cellular carbon ranged between 8.3 to 79.4% of the total picophytoplankton (<3 m), and between 4.7 to 46.4% of the total phytoplankton. The larger percentages of the picocyanobacteria occurred consistently in low chlorophyll waters.  相似文献   

20.
The relationship between life-history characteristics and population dynamics were investigated in the onuphid polychaeteKinbergonuphis simoni (Santos, Day and Rice) between 1982 and 1987. The studied population is located in Upper Tampa Bay, Florida, USA. This worm attains sexual maturity at 5 to 10 mo of age, depending on temperature. Several consecutive broods are produced during a female's life time, with 7 to 26 young per brood. Adults die in June–July after the breeding season. Generations of breeding individuals do not overlap and the replacement of generations occurs in July–August. Only one extended breeding season is experienced per life time. Life span does not exceed 2 yr. Field population density in 1982 was high in fall and spring (2000 to 9000 individuals/m2), low in summer (1600 individuals/m2), and slightly depressed in winter (2500 to 4000 individuals/m2). Changes in population density may be explained by seasonal, temperature-controlled changes in instantaneous birth and death rates. Birth rates are high in fall and spring, while death rates are high in early summer. Death of juveniles contributes to the winter density decrease.  相似文献   

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