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1.
Carotenoids are the most universal and most widespread pigments in nature. They have played pivotal roles in the evolution of photosensing mechanisms in microbes and of vision in animals. Several groups of phytoflagellates developed a photoreceptive organelle called the eyespot apparatus (EA) consisting of two separable components: the eyespot, a cluster of carotenoid‐rich globules that acts as a reflector device, and actual photoreceptors for photobehaviors. Unlike other algal eyespots, the eyespot of Euglenophyta lacks reflective properties and is generally considered to act as a shading device for the photoreceptor (paraflagellar body, PFB) for major photomovements. However, the function of the eyespot of Euglenophyta has not yet been fully proven. Here, we report that the blocking carotenoid biosynthesis in Euglena gracilis by suppressing the phytoene synthase gene (crtB) caused a defect in eyespot function resulting in a loss of phototaxis. Raman spectroscopy and transmission electron microscopy suggested that EgcrtB‐suppressed cells formed eyespot globules but had a defect in the accumulation of carotenoids in those packets. Motion analysis revealed the loss of phototaxis in EgcrtB‐suppressed cells: a defect in the initiation of turning movements immediately after a change in light direction, rather than a defect in the termination of cell turning at the appropriate position due to a loss of the shading effect on the PFB. This study revealed that carotenoids are essential for light perception by the EA for the initiation of phototactic movement by E. gracilis, suggesting one possible photosensory role of carotenoids in the EA for the phototaxis.  相似文献   

2.
Summary A comparative action spectroscopical study was made on phototaxis in two genera of cryptomonads (cryptophyte flagellate algae), namely,Cryptomonas (rostratiformis) andChroomonas (nordstedtii andcoeruled). The two genera differ in their characteristic phycobilin pigmentation and, among three species, onlyChroomonas coerulea possesses an eyespot. The two species with no eyespot,Cryptomonas rostratiformis andChroomonas nordstedtii, exhibited positive phototaxis, showing very similar action spectra characterized by a broad band in the region from 450 nm to 650 nm, with an action maximum at about 560 nm; these features are essentially the same as those observed previously forCryptomonas strain CR-1. InCryptomonas rostratiformis, a small peak was also found at 280 nm in the UV-B/C region.Chroomonas coerulea, with eyespot, did not exhibit distinct positive phototaxis in a wide spectral region at any given, even very low, light intensity, but exhibited negative phototaxis of spectral sensitivity maximal at 400–450 nm. These results indicate that the positive phototaxis ofCryptomonas (rostratiformis and CR-1) andChroomonas nordstedtii is mediated by the same, yet unidentified photoreceptor(s).Chroomonas nordstedtii, possessing no phycoerythrin absorbing at 545 nm, also exhibits positive phototaxis at ca. 560 nm, and this result disfavors the so far proposed possibility that the positive phototaxis of the cryptophytes may be mediated by phycobilin pigments. On the other hand, the spectral characteristics of negative phototaxis ofChroomonas coerulea can possibly be ascribed to the presence of an eyespot.  相似文献   

3.
Chlamydomonas reinhardtii exhibits photophobic and positive and negative phototactic responses that can be defined for cell populations using computerized cell tracking and motion analysis. Mutants CC-2359 and FN68 are pigment deficient mutants that are blocked in carotenoid synthesis and lack these photo responses. In particular, neither mutant exhibits flash-induced photophobic responses to visible light stimuli to which wild-type gametic cells exhibit a strong response, with several behavioral stages. Upon addition of all-trans retinal to these mutants, the photophobic responses are restored with minor quantitative differences from wild-type populations. Using both light and electron microscopy, we have compared the ultrastructural characteristics of wild-type C. reinhardtii to those of both mutants. As previously described, wild-type cells contain an eyespot consisting of 2–4 layers of pigmented granules encased within thylakoid membranes, located between the distal extremities of the flagellar root. This structure is also visible as an orange-red spot in light microscopy. The photoreceptor is thought to be concentrated in the plasma membrane above the eyespot. The mutant, CC-2359, lacks this eyespot as seen by both light and electron microscopy, even when the photophobic response has been restored. FN68-like mutants studied earlier by Morel-Laurens and Feinlieb and others contain an eyespot which can be seen only by electron microscopy. In FN-68, the eyespot generally has the same dimensions as in wt cells, differing mainly in pigment granule appearance. Consistent with these findings, several laboratories have shown that the full range of phototactic responses can be reconstituted in FN68 and CC-2359, but that negative phototaxis requires a significantly stronger light stimulus in the latter strain. We confirm the suggestion that the eyespot is not necessary for the photophobic response, and is not critical for the appropriate assembly and function of the photophobic response receptor in the membrane. Furthermore, the locus of reconstitution of the functional receptor is not the eyespot. Because of the definitive demonstration of the absence of the eyespot in CC-2359, however, the eyespot may play a role in negative phototaxis.  相似文献   

4.
Great structural variety is seen in the eyespot of dinoflagellates, a structure involved in phototaxis. Although there are several works on the phototactic responses in some species of dinoflagellates, none of the dinoflagellates used in these studies possessed an eyespot and, therefore, we have no knowledge of the relationship between eyespot type and phototactic response. In this study, we determined wavelength dependency curves for phototaxis in four marine dinoflagellates that possess a different type of either eyespot or chloroplast. These include: (i) a dinoflagellate possessing a peridinin-containing ohioroplast with an eyespot (Scrippsiella hexapraecingula Horiguchi et Chihara); (ii) a dinoflagellate containing a diatom endosymbiont and with the type B eyespot sensu Dodge (1984; (Peridinium foli-aceum (Stein) Biecheler); (iii) a dinoflagellate with peri-dinin-containing chloroplasts, but lacking an eyespot (Atexandrium hiranoi Kita et Fukuyo); and (iv) a dinoflagellate with fucoxanthin, 19′-hexanoyloxyfucoxanthin and 19′-butanoyloxyfucoxanthin, but lacking an eyespot (Gymnodinium mikimotoi Miyabe et Kominami ex Oda), Regardless of the eyespot or the chloroplast type, all four dinoflagellates showed similar wavelength dependency curves for phototaxis, with sensitivity between 380 and 520 nm, the highest peak at approximately 440 or 460 nm and smaller peaks or shoulders at 400–420 nm and 480–500 nm. Substantial peaks have also been noted in the ultraviolet range (260–280 nm). The ultrastructural study of the eye-spot of Scrippsiella hexapraecingula revealed that the eyespot consists of two layers of lipid globules and probably acts as a quarter-wave stack antenna.  相似文献   

5.
M. Melkonian  H. Robenek 《Protoplasma》1979,100(2):183-197
Summary The eyespot region of the flagellateTetraselmis cordiformis Stein (Chlorophyceae) was investigated with the freeze-fracture technique. The only fracture faces observed in this region were the two complementary fracture faces (PF and EF) of the outer chloroplast envelope membrane. Intramembranous particle numbers on both fracture faces of this membrane were significantly higher in the eyespot region as compared to regions outside the eye-spot. Higher numbers of particles on the PF face in the eyespot region were mainly caused by an increase in particle numbers of the size class 6–8 mm, while on the EF face particle size distribution was not significantly different between eyespot and other regions. Functional implications are discussed and evidence is presented that the outer chloroplast envelope membrane may be the site of photoreceptor location in green algal phototaxis.  相似文献   

6.
The eyespot of Chlamydomonas reinhardtii is a light-sensitive organelle important for phototactic orientation of the alga. Here, we found that eyespot size is strain specific and downregulated in light. In a strain in which the blue light photoreceptor phototropin was deleted by homologous recombination, the light regulation of the eyespot size was affected. We restored this dysfunction in different phototropin complementation experiments. Complementation with the phototropin kinase fragment reduced the eyespot size, independent of light. Interestingly, overexpression of the N-terminal light, oxygen or voltage sensing domains (LOV1+LOV2) alone also affected eyespot size and phototaxis, suggesting that aside from activation of the kinase domain, they fulfill an independent signaling function in the cell. Moreover, phototropin is involved in adjusting the level of channelrhodopsin-1, the dominant primary receptor for phototaxis within the eyespot. Both the level of channelrhodopsin-1 at the onset of illumination and its steady state level during the light period are downregulated by phototropin, whereas the level of channelrhodopsin-2 is not significantly altered. Furthermore, a light intensity–dependent formation of a C-terminal truncated phototropin form was observed. We propose that phototropin is a light regulator of phototaxis that desensitizes the eyespot when blue light intensities increase.  相似文献   

7.
  1. Recent evidence suggests that periphytic algae stimulate plant litter heterotrophs (fungi and bacteria) in the presence of light, but few studies have tested whether this stimulation varies across gradients of light, which may covary with temperature.
  2. We exposed field-conditioned Typha domingensis litter to fully-crossed, short-term gradients of temperature (15, 20, 25, and 30°C) and light (0, 25, 53, 123, and 388 µmol quanta m−2 s−1) and measured responses of litter-associated algal, fungal, and bacterial production rates and β-glucosidase, β-xylosidase, and phenol oxidase enzyme activities in the laboratory.
  3. Increased light stimulated algal production rates, from immeasurable production under darkness to >200 µg algal C g−1 detrital C hr−1 at the highest light level, with the greatest light sensitivity and maximal photosynthetic rates at 25°C. In turn, increased light stimulated fungal production rates, especially at the two highest temperatures and most strongly at 25°C where light stimulated fungal production by a mean of 65 µg C g−1 detrital C hr−1, indicating 2.1-fold stimulation by light. Bacterial production rates also responded to light, indicated by stimulation of a mean of 16 µg C g−1 detrital C hr−1 (1.6-fold) at 15°C, but stimulation was weaker at higher temperatures. Enzyme activities increased strongly with elevated temperature but were not affected by light.
  4. Our experimental evidence suggests algae differentially stimulate litter-associated bacteria and fungi in a light-dependent manner that further depends on temperature. These findings advance understanding of the onset of algal stimulation of heterotrophy, including algal-induced priming effects during litter decomposition, in response to common covarying environmental gradients subject to global change.
  相似文献   

8.
Lifetable demography and reproductive traits of a Kenyan strain of the rotifer Brachionus angularis were investigated using individual and small batch culture approaches. The rotifer was identified morphologically before conducting studies at 20, 25 and 30 °C, using Chlorella vulgaris at 2.5 × 105 to 2.5 × 107 cells ml–1. The rotifers were highly fecund, producing 2.11 ± 0.07 offspring female–1 day–1 and reproductive, producing 8.43 ± 0.24 offspring female–1 at 25 °C with 2.5 × 106 algal cells ml–1. The highest intrinsic rate of natural increase (0.74 ± 0.02 d–1), specific population growth rate (0.49 ± 0.01), longest life expectancy at hatching (12.41 ± 0.28 d) and shortest generation time (2.87 ± 0.03 d) also occurred at 25 °C with 2.5 × 106 algal cells ml–1. The duration of hatching to first spawning was shortest (2.86 ± 0.21 h) at 30 °C with 2.5 × 107 algal cells ml–1 and longest (8.83 ± 0.39 h) at 20 °C with 2.5 × 105 algal cells ml–1. The highest population density (255.7 ± 12.6 ind. ml–1) was realised at 25 °C with 2.5 × 106 cells ml–1 on Day 8, whereas the lowest population density (122.0 ± 3.6 ind. ml–1) was realised at 20 °C with 2.5 × 105 cells ml–1 on Day 8. The lorica length and width of the Kenyan strain of B. angularis are 85.6 ± 3.1 µm and 75.4 ± 3.6 µm, respectively. The rotifer optimally reproduces at 25 °C when fed with 2.5 × 106 algal cells ml–1.  相似文献   

9.
The fine structure of the binucleate, fucoxanthin-containing dinoflagellate Peridinium foliaceum (Stein) Biechler was re-examined for evidence of an endosymbiout. The eucaryotic nucleus, chloroplasts and associated ribosome-dense cytoplasm were separated by a single invaginating membrane from the rest of the dinoflagellate cell. The triple membrane-enclosed eyespot, mesocaryotic nucleus, trichocysts and accumulation bodies resided in the dinoflagellate cytoplasm. These observations suggest that P. foliaceum contains a membrane-bound endosymbiont, similar to that already described for the closely related species. P. balticum (Levander) Lemmermann.  相似文献   

10.
The function of the eyespot in phototaxis of the flagellate green alga Chlamydomonas reinhardtii Dangeard was studied using quantitative reflection confocal laser scanning microscopy and photoelectric measurements. The reflective properties of the eyespot and the photoreceptor current of the C. reinhardtii eyespot mutant ey 627, mt were compared with those of Chlamydomonas strains possessing a well-developed eyespot. Under growth conditions in which strongly disorganized eyespots were observed in the mutant by electron microscopy, there was a significant reduction in the reflection intensity of the eyespot and in the amplitude ratio (500440 nm) of photoreceptor currents induced by flashes of 500- and 440-nm light in non-oriented cells. Photoelectrical responses of pre-oriented cells revealed that the latter effect is caused by an altered directional sensitivity of the antenna complex, whereas the functional state of the photoreceptor pigment is not strongly affected in mutant cells. Both the reflection intensity and the amplitude ratio of photoreceptor currents increased to the level of reference strains under conditions supporting the development of a well-organized eyespot in the mutant. Furthermore, incubation of the mutant with high concentrations of all-trans-retinal (10 M), independent of whether carotenoid biosynthesis was inhibited or not, was found to increase the reflection intensity of the eyespot. An increase in the rate of photoorientation of the mutant occurred concomitant with the increase in the reflective properties of the mutant eyespot. These observations demonstrate the importance of an intact eyespot for interference reflection and absorption of phototactically active light, and thus for the directional sensitivity of the eyespot apparatus.Abbreviations HSM high-salt medium This study was supported by the Deutsche Forschungsgemeinschaft. O. A. Sineshchekov was supported by a Research Fellowship from the Alexander von Humboldt Foundation. The authors wish to thank U. Powalowski (Botanisches Institut, Universität zu Köln) for help with electron microscopy.  相似文献   

11.
We have isolated a new Chlamydomonas reinhardtii Dangeard (Chlamydomonadales, Chlorophyceae) mutant with from one up to more than four eyespots cell?1. It was designated mes (multiple eyespots)‐10 A wild‐type cell has a single eyespot, located under the chloroplast envelope, at a certain position near the cell's equator where the chloroplast envelope is in contact with the cell membrane. The eyespot(s) in mes‐10, however, are located at various positions on its chloroplast. The mes‐10 cells displayed negative phototaxis to 480–500 nm light. This behavior differed from that of a similar mutant, ptx4, which has been shown to have multiple eyespots and display no phototaxis (Pazour et al., J. Cell Biol. 1995; 131 : 427–40). Mes‐10 may retain a functional photoreceptor and a photosignal transduction system independently of its multiple eyespots. This mutant should be useful for studying how C. reinhardtii responds to light signals, as well as how eyespots are formed in the cell.  相似文献   

12.
Growth of Pseudocercosporella herpotrichoides on potato dextrose agar at water potentials from -0.5 to -6.9 MPa was optimal at 20°C. At 12 and 20°C, six isolates of P. herpotrichoides grew more rapidly at -0.5 to -2 MPa than at -6.9 MPa. Wheat plants inoculated with P. herpotrichoides and grown in columns of soil at either 15 or 20°C developed more severe eyespot lesions under a heavy watering regime than under medium or light watering regimes. P. herpotrichoides penetrated leaf sheaths of inoculated plants grown in compost more rapidly at 10°C night/15°C day temperatures than at 5/10°C; death of leaf sheaths was also more rapid and consequently there were fewer living infected leaf sheaths at 10/15°C than at 5/10°C. Irrigating for 5 wk before harvest increased the severity of eyespot lesions in a 1983 wheat crop.  相似文献   

13.
A. McMinn  A. Martin  K. Ryan 《Polar Biology》2010,33(11):1547-1556
The phytoplankton and sea ice algal communities at the end of winter in McMurdo Sound were dominated by Fragilariopsis sublineata, with Thalassiosira antarctica, Melosira adele, Pinnularia quadreata, Entomoneis kjellmannii and heterotrophic dinoflagellates also present. Sea ice algal biomass at the end of winter was very low, only 0.050 ± 0.019 mg chla m−2 in 2007 and 0.234 ± 0.036 mg chla m−2 in 2008, but this increased to 0.377 ± 0.078 mg chla m−2 by early October in 2007 and to 1.07 ± 0.192 by late September in 2008. Under ice phytoplankton biomass remained consistently below 0.1 μg chla l−1 throughout the measuring period in both years. The photosynthetic parameters Fv/Fm, rETRmax and α document microalgal communities that are mostly healthy and well adapted to their low light under ice environment. Our results also suggest that species such as Fragilariopsis sublineata are well adapted to deal with low winter light levels but are unlikely to survive an increase in irradiance, whereas other taxa, such as Thalassiosira antarctica, will do better in a higher light environment.  相似文献   

14.
A new sand‐dwelling dinoflagellate from Palau, Galeidinium rugatum Tamura et Horiguchi gen. et sp. nov., is described. The life cycle of this new alga consists of a dominant nonmotile phase and a brief motile phase. The motile cell transforms itself directly into the nonmotile cell after swimming for a short period, and cell division takes place in the nonmotile phase. The nonmotile cell possesses a dome‐like cell covering, which is wrinkled and equipped with a transverse groove on the surface. The cell has 10–20 chloroplasts and a distinct eyespot. The motile cell is Gymnodinium‐like in shape. The dinoflagellate possesses an endosymbiotic alga to which the chloroplasts belong and which is separated from the host (dinoflagellate) cytoplasm by a unit membrane. The endosymbiont cytoplasm also possesses its own eukaryotic nucleus and mitochondria. The eyespot is surrounded by triple membranes and is located in the host cytoplasm. Photosynthetic pigment analysis, using HPLC, revealed that G. rugatum possesses fucoxanthin as the principal accessory pigment instead of peridinin. The rbcL tree showed that G. rugatum is monophyletic with Durinskia baltica (Levander) Carty et Cox and Kryptoperidinium foliaceum (Stein) Lindemann and that this clade is closely related to the pennate diatom, Cylindrotheca sp. The endosymbiont of G. rugatum is therefore shown to be a diatom. Phylogenetic analysis based on small subunit rDNA sequences demonstrated that G. rugatum, D. baltica, and K. foliaceum, all of which are known to harbor an endosymbiont of diatom origin, are closely related.  相似文献   

15.
Flagellate green algae have developed a visual system, the eyespot apparatus, which allows the cell to phototax. To further understand the molecular organization of the eyespot apparatus and the phototactic movement that is controlled by light and the circadian clock, a detailed understanding of all components of the eyespot apparatus is needed. We developed a procedure to purify the eyespot apparatus from the green model alga Chlamydomonas reinhardtii. Its proteomic analysis resulted in the identification of 202 different proteins with at least two different peptides (984 in total). These data provide new insights into structural components of the eyespot apparatus, photoreceptors, retina(l)-related proteins, members of putative signaling pathways for phototaxis and chemotaxis, and metabolic pathways within an algal visual system. In addition, we have performed a functional analysis of one of the identified putative components of the phototactic signaling pathway, casein kinase 1 (CK1). CK1 is also present in the flagella and thus is a promising candidate for controlling behavioral responses to light. We demonstrate that silencing CK1 by RNA interference reduces its level in both flagella and eyespot. In addition, we show that silencing of CK1 results in severe disturbances in hatching, flagellum formation, and circadian control of phototaxis.  相似文献   

16.
Patterns of larval release, dispersal and settlement in sponges are poorly understood despite their significance in explaining adult ecology. Time of release, swimming speeds, phototaxis and vertical migration were quantified for larvae of the dictyoceratid sponge Coscinoderma matthewsi. The influence of cues associated with biofilms and coral rubble on larval settlement and metamorphosis was also measured. C. matthewsi is a brooding sponge and releases tufted parenchymellae larvae during the day. Upon release, larvae (>90%) have no phototactic response, maintaining their position at the water surface for 80 min ± 0 (mean ± SE) regardless of a light cue (natural daylight) before exhibiting negative phototaxis. At 28 h post-release, the majority of larvae (94.7% ± 6.1) exposed to light from the surface migrated to the bottom and assumed a demersal phase. Without light, larvae occupied the surface for up to 28 h post-release (89.3% ± 1.8) before migrating to the bottom. Larvae did not settle gregariously and began to settle and metamorphose after 28 h post-release without a cue. Settlement and metamorphosis were faster in the presence of a biofilm (settlement = 15.0% ± 8.7 and metamorphosis = 12.5% ± 9.5 at 28 h post-release), while the addition of coral rubble accelerated metamorphosis further (settlement = 10.0% ± 4.1 and metamorphosis = 27.5% ± 10.3 at 28 h post-release) compared to controls (sterile surfaces) (settlement = 0% and metamorphosis = 0% at 28 h post-release). However, both biofilms and coral rubble decrease total metamorphosis (control = 92.5% ± 4.8, biofilms = 67.5% ± 7.5 and coral rubble = 55.0% ± 13.2) due to mortality after 76 h post-release.  相似文献   

17.
SYNOPSIS. The eyespot of the zoospore of Tetracystis excentrica (a green alga) has been studied by light and electron microscopy. In Tetracystis the eyespot consists of about 110 osmiophilic granules which form a plate in the anterior third of the cell. The granules are about 80 Å in diameter and are found in the outermost portion of the chloroplast; they commonly show hexagonal close packing and a hexagonal shape. The granules are confined positionally by the chloroplast envelope and an inner thylakoid. The plasmalemma over the eyespot is thickened and is separated from the chloroplast envelope by a 50 mμ space. The eyespot of Tetracystis is compared with others reported in the literature and the possible functional significance of these studies is discussed. The possibility that the eyespot plate in Tetracystis serves as a shading device rather than the primary photoreceptor is considered.  相似文献   

18.
Behavior of the eyespots during the fertilization of Ulva arasakii Chihara was studied using field emission scanning electron microscopy (FE‐SEM). FE‐SEM enabled the visualization of the eyespot of biflagellate male and female gametes. The smaller male gamete has one protruded smaller (1.3 ± 0.15 μm× 1.0 ± 0.29 μm) eyespot and the larger female gamete has a larger (1.6 ± 0.2 μm× 1.1 ± 0.13 μm) one on a posterior position of the cell. The cell membrane over the eyespot region is relatively smooth compared to other parts of the cell body and exhibits hexagonal arranged lipid globules. Because the size of the cell and the morphology of the eyespot are different between male and female gametes, we could follow the fate of the eyespots during the fertilization. The initial cytoplasmic contact and fusion of the gametes takes place at their anterior end, slightly posterior to the flagellar base. The morphology of the fusing gametes followed two clearly distinguishable patterns. About half the gamete pairs lie side‐by‐side with their longitudinal axes nearly parallel, while the rest are oriented anti‐parallel to each other. In all cases, the larger female gamete fused along the same side as the eyespot, while the smaller male gamete fused along the side away from its eyespot. As fusion proceeds, the gamete pair is transformed into the quadriflagellate planozygote, in which the eyespots are positioned side‐by‐side on the region of cell fusion. These observations indicated that the opposite positioning of the eyespot relative to the cell fusion site in male and female gametes is important for the proper arrangement of the eyespots in the planozygote. The significance of this feature in advanced green algae is briefly discussed.  相似文献   

19.
We describe three coccoid green algal strains belonging to a new genus and species, Marinichlorella kaistiae Z. Aslam, W. Shin, M. K. Kim, W.‐T. Im et S.‐T. Lee, in seawater samples from the South Sea of Korea. These strains were maintained at 25°C–30°C under a 12:12 light:dark (L:D) photoregime in an ASN‐III medium at a pH of 7.5. These strains were tolerant of high salinity (7.5% NaCl) (w/v) and temperature (40°C). Molecular phylogenetic analyses using 18S rRNA gene sequence data resolved these organisms to a clade separate from green coccoid algae with similar morphology. The DNA–DNA hybridization results demonstrated very low relatedness of these organisms to phylogenetically related species of the genera Chlorella and Parachlorella. The molar guanine + cytosine content (G + C mol%) of the genomic DNA of these organisms ranged from 64.7 to 69.1 mol%. Based on molecular phylogeny, DNA–DNA hybridization, and other morphological studies, we propose a new taxon, Marinichlorella kaistiae, to describe these strains and classify them in the family Chlorellaceae. The type strain is KAS007T (= KCTC AG10303T = IAM C‐620T).  相似文献   

20.
Laboratory assays demonstrated the presence of a small positive geotaxis response to a 15° incline by Folsomia candida Willem (Collembola: Isotomidae). Negative phototaxis played an additive role to positive geotaxis when the experimental apparatus were exposed to light. The geotactic response was negatively affected by cold acclimation and decreasing surrounding temperature, but unaffected by food deprivation. The reduced mobility of springtails at low temperature did not seem to play a role in the corresponding decreased geotaxis. The low level of geotaxis and its further decrease with exposure to low temperature support an earlier suggestion that F. candida do not respond to cooling temperatures of fall by relocation to warmer deeper soil layers, but remain in the upper soil layers and increase their cold tolerance to continue foraging in the food‐rich upper soil layers.  相似文献   

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