Effect of aluminum on callose synthesis in root tips of tea (Camellia sinensis L.) plants

Aluminum (Al) toxicity is a major factor limiting yield production on acid soils (Foy 1983). The initial symptom of Al toxicity in many plants is manifested by the inhibition of root elongation (Ownby and Popham 1990; Llugany et al. 1994; Sasaki et al. 1994; Horst et al. 1997), which occurs during a very short period of time after exposure to Al (Llugany et al. 1994; Staß and Horst 1995). In a large number of recent reports, it was shown that the root apex plays a major role in the Al-sensitivity and response mechanisms (Zhang et al. 1994; Sasaki et al. 1997; Sivaguru and Horst 1998). However, it is interesting to note that stimulatory effects of Al on the growth of plants have also been reported in some studies (Chenery 1955; Konishi et al. 1985; Huang and Bachelard 1993; Osaki et al. 1997). In tea plant (Camellia sinensis L.) a stimulatory effect of Al on the growth was also demonstrated in some experiments, using intact plant (Chenery 1955; Konishi et al. 1985), cultured roots (Tsuji et al. 1994), and pollen tubes (Yokota et al. 1997). The growth of tea roots was typically more stimulated than that of shoots by Al (Konishi et al. 1985). It was assumed that Al effects might be due to the amelioration of phosphorus absorption (Konishi et al. 1985), secretion of malic acid from roots to dissolve aluminum phosphate in the rhizosphere (Jayman and Sivasubramaniam 1975), stimulation of growth of microorganisms on the root surface (Konishi 1990) or replacement of some functions of boron (Konishi 1992; Yokota et al. 1997). However, the stimulatory effects of Al on tea plant growth have not yet been el ucidated.

The formation of callose (1,3-β-glucan) has been reported as a common plant response to a variety of stresses, as well as mechanical, biophysical, chemical, and biological injury (Jaffe and Leopold 1984; Zhang et al. 1994). Increased synthesis of callose has been observed upon exposure to excess amounts of some elements, such as boron (McNairn and Currier 1965), cobalt, nickel, zinc (Peterson and Rauser 1979), and manganese (Wissemeier and Horst} 1987, 1992). Callose synthesis was also induced by Al in the roots of Triticum aestivum (Zhang et al. 1994) and Zea mays (Horst et al. 1997; Sivaguru and Horst 1998), suspension-cultured cells of Glycine max (Staß and Horst 1995), and protoplasts of Avena sativa (Schaeffer and Walton 1990) and Zea mays (Wagatsuma et al. 1995). Induction of callose synthesis in roots seems to be a very rapid physiological indicator of Al-induced injury or genotypical differences in Al sensitivity (Wissemeier and Horst 1992; Zhang et al. 1994; Horst et al. 1997). Nevertheless, Al-induced callose synthesis in tea plant, whose growth is stimulated by suitable Al concentrations, has not been described yet. Therefore, to elucidate the physiological basic effects of Al on tea plant, callose synthesis affected by Al in the root tips of intact plants was analyzed in the present study.     

Ontogeny and total sugar content of yacon tuberous roots and other three Smallanthus species (Heliantheae,Asteraceae), insights on the development of a semi-domesticated crop

The ontogeny of the tuberous roots of yacon (Smallanthus sonchifolius), an ancient Andean crop, is described for the first time along with three other wild species of Smallanthus: S. connatus, S. macroscyphus and S. siegesbeckius. Tuberous storage roots are present in the four species. Tuberization results from unusual secondary thickening. A meristematic endodermis maintains the primary cortex by generating cortex cells continuously. Secretory schizogenous canals develop adjacent to the endodermis in opposition to the phloem poles. Phellogen originates in the outer layers of the parenchymatic cortex. The four species develop a parenchymatous pith that stores mainly fructooligosacharides (FOS) and plays an important role in storing water, protecting the plant from drought stress and environmental hazards in the Andean climate. In yacon, domestication has produced a wide pith that stores water, reducing sugars and FOS with low degree of polimerization, that could have eventually made it more palatable as compared to the wild counterparts.     

Shoot control of nodulation is modified by the root in the supernodulating soybean mutant En6500 and its wild-type parent cultivar Enrei

Abstract

Based on grafting studies, both supernodulating (Carroll et al. 1985a, b) and hypernodulating (Gremaud and Harper 1989) soybean ([itGlycine max} L. Merr.) phenotypes were reported to be under the control of shoot factors (Delves et al. 1986, 1987; Cho and Harper 1991). Recently Akao and Kouchi (1992) have isolated a new supernodulating mutant (En6500) from ethyl methane sulfonate (EMS)-treated Enrei, a cultivar which is widely grown in the central districts of Japan. This mutant has been shown to produce several fold as many nodules as its wild-type parent cultivar when grown at a low concentration (0.5 mol m^-3) of nitrate. Moreover, it exhibited a continuous increase in the nodule number with the increasing nitrate concentration, even at 15 mol m^-3 (Francisco et al. 1992), in contrast to the similar mutant nts382 in which the nodulation decreased even at the relatively low nitrate level of 5.5 mol m^-3 (Carroll et al. 1985a). In this study we conducted grafting experiments to determine which plant part controls the supernodulation of En6500.     

Optimum Nutrient Level on Growth,Flowering, and Rhizome Production in Curcuma

ABSTRACT

This study investigated the effect of liquid fertilizer treatments on growth, flowering, leaf mineral content, and rhizome production during forcing of Curcuma alismatifolia ‘Chiang Mai Pink’ and C. thorelii ‘Chiang Mai Snow’. Plants were irrigated with 200 mL of 1.3 g L^?1 of 15 nitrogen (N) -7 phosphorus (P) -14 potassium (K) water soluble fertilizer at 0, 1.3, 2.7, 4.0, 5.3, or 6.6 g L^?1 weekly. Days to flower, flower stem length, and leaf length were recorded, the mineral contents in leaves were analyzed, and the number of rhizomes with tuberous roots were recorded at harvest. Flowering of the first inflorescence in both C. alismatifolia ‘Chiang Mai Pink’ and C. thorelii ‘Chiang Mai Snow’ was significantly delayed when plants received 6.6 g L^?1 fertilizer as compared to the control plants. The number of rhizomes with more than 4 tuberous roots was highest when plants received 2.7 g L^?1 fertilizer. No medium-sized rhizomes with more than seven tuberous roots were produced when ‘Chiang Mai Pink’ plants received 0, 4.0, 5.3, and 6.6 g L^?1 fertilizer. Based on the production of rhizomes with four to six tuberous roots, optimum concentration of 15N -7P -14K water soluble fertilizer is 2.7 g L^?1 for C. alismatifolia ‘Chiang Mai Pink’ and 1.3 to 4.0 g L^?1 for C. thorelii ‘Chiang Mai Snow’. Although high boron content occurred only in the outer part of the second leaf when fertilizer concentrations were increased, leaf-margin burn (LMB) symptoms were not observed in both species and this could not be related to the production of rhizomes.     

The biological and chemical variability of yacon

This paper focuses on the biological and chemical variability of four yacon (Smallanthus sonchifolius) accessions cultivated under field conditions. Significant variations in tuber shape, weight, content of oligofructans, as well as in leaf isozymes, phenolics, and relative DNA contents were found. Accessions 6 and 88 were the most productive (up to 3.01 and 3.74 kg/plant); accession 48 was the most balanced from the yield aspect in three vegetative periods. A significantly higher content of beta-(2-->1) oligofructans was noted in accessions 48 and 88 as compared to 6 and 60. No difference in sucrose, glucose, and fructose level was observed. Only accession 6 exhibited separate acid phosphatase and esterase isoforms. Accessions 6 and 60 had the highest content of phenolics, and accession 88 had the lowest relative DNA content. Large yacon intraspecific variation may be useful in future detailed research as a good background for breeding, growing, and utilization in industrial processing.     

Sulfur deficiency of rice plants in the Lower Volta area,Ghana

Abstract

It is well known that stem nodules are formed on the aerial parts of Aeschynomene spp. and Sesbania rostrata grown in the field (Yatazawa and Yoshida 1979; Dreyfus and Dommergues 1981; Yoshida et al. 1985). We have reported that stem nodules were successfully formed by inoculation of Rhizobium isolates derived from both stem and root nodules of A. indica (Yoshida et al. 1985; Sasakawa et al. 1986). The specific activity of nitrogen fixation in stem nodules is comparable to that of root nodules (Sasakawa et al. 1986; Sasakawa 1990). A red pigment, which suggests the presence of leghemoglobin, was detected in stem nodules as well as in root nodules (Yatazawa and Yoshida 1979; Yatazawa and Susilo 1980; Sasakawa et al. 1986).     

Effect of micronutrient application on the growth and occurrence of sterility in barley and rice in a Malaysian deep peat soil

Abstract

Nodule formation in legume crops is a multistep process which involves the interactive gene expression of bacteria and host plants. Attempts to widen the genetic variation of symbiotic bacteria and host plants have led to the isolation of several mutants with a diverse capability of nodulation. The most striking variants in hosts are the supernodulating mutants isolated in pea (Pisum sativum L.) (Jacobsen and Feenstra 1984; Duc and Messager 1989), common bean (Phaseolus vulgaris L.) (Park and Buttery 1988), and soybean (Glycine max (L.) Men.) (Carroll et al. 1985a,b; Gremaud and Harper 1989; Akao and Kouchi 1992), all of which are capable of producing several-fold more nodules than their parental lines in the presence of nitrate. These mutants may be useful materials for analyzing the mechanism controlling nodule formation, and are considered to have a high agronomic potential under certain growing conditions.     

Characterization of Desulfovibrio sp. isolated from some lowland paddy field soils of Burkina Faso

In a wetland ecosystem such as lowland ricefields, the anaerobic mineralization of organic matter is a key mechanism for nutrient recycling (Jørgensen 1982; Freney et al. 1982). In the process which involves several bacterial groups, sulfate reducers (Watanabe and Furusaka 1980; Widdel 1988; Ouattara and Jacq 1992; Nozoe 1997), methanogens (Asakawa and Hayano 1995; Dianou et al. 1997), sulfur and ferric ion reducers (Jacq et al. 1991) become active when the soil becomes anoxic (Amstrong 1969). Sulfate reducers are common in flooded soils (Watanabe and Furusaka 1980; Widdel 1988; Furusaka et al. 1991). Reported densities in rice soils ranged from 10³ to 10⁵ g^-1 dry soils in Asia (Watanabe and Furusaka 1980) and from 10² to 10⁹ g^-1 dry soil in Sénégal (Ouattara and Jacq 1992). Among the sulfate reducers, the “classical” Desulfovibrio species are known to oxidize typical fermentation products such as hydrogen, lactate, pyruvate, and dicarboxylic acids (Widdel 1988). So far, data on the isolation and characterization of sulfate-reducing bacterial (SRB) strains from African rice paddy field soils study have been very limited.     

Occurrence of 4-methyleneglutamine and 2-oxo-4-methyl-3-pentene-1,5-dioic acid in leaves and stem of tulip plants

Abstract

The presence of 4-MeGln in the tulip plant was discovered by ZACHARIUS et al. (1954), and it was found that this amide generally occurred in the leaves of almost all the species of genus Tulipa (FOWDEN and STEWARD 1957a). The 4-MeGln compound has been detected in every part of the tulip plant, i.e., bulb scales, roots, basal plate, young shoots, leaves, stern, and flower (FOWDEN and STEWARD 1957a, b; OHYAMA 1986; OHYAMA et al. 1985, 1988a, b; ZACHARIUS et al. 1954, 1957). Especially 4-MeGln was found to be a major soluble N constituent in the leaves and stem of tulip of the flowering stage (OHYAMA et al. 1985; OHYAMA 1986).     

Effect of root systems on preferential flow in swelling soil

Abstract

Permeability problems on irrigated soils may be alleviated by root systems that increase water flow by creating macropores. Infiltration rates have been shown to increase where plant roots decay and serve as preferential flow paths. For low‐organic‐matter swelling soil, there is a question whether macropores are able to resist the lateral swelling forces of the soil. The objective of this study was to observe preferential water flow paths in a swelling soil under two cropping systems. A Holtville silty clay (clayey‐over‐loamy, montmorillonitic Typic Torrifluvent) was observed in situ. Two crops, alfalfa (Medicago sativa, L.) and wheat (Triticum turgidum, L.) provided sharply contrasting root systems, with wheat possessing fine, fibrous roots; alfalfa on the other hand, has a taproot system. Macropores were observed after applying soil‐adsorbing methylene blue dye to irrigation water. Shrinkage cracks failed to conduct dye after 10 minutes into a flood irrigation. Earthworm (Lubricus terrestris) channels were also not stable. However, decaying roots of alfalfa produced stable macropores, while wheat produced no such macropores. The influence of alfalfa‐root‐induced macropores was demonstrated by the increase in final infiltration rate during alfalfa cropping which agreed with Meek et al.’s (1989, 1990) findings on sandy loam soils.     

Amino acid sequence of proteins induced in Fe-deficient stressed alfalfa (Medicago sativa L.) roots

Alfalfa (Medicago sativa L.) grows well in soils with a moderately high pH and dissolves insoluble iron in the rhizosphere. We have investigated active uptake mechanisms under Fe-deficient nutrient conditions and the effects of Fe-deficiency on plants. Previously, we observed that Fe-deficient alfalfa roots exuded many compounds (Masaoka et al. 1993) such as fiavonoids. We also identified a new compound “alfafuran” which is a phenol compound and is different from organic acids or phytosiderophore-type amino acid derivatives exuded by Fe-deficient plant roots. This compound is also very effective in dissolving ferric phosphate (Noguchi et al. 1994), suggesting that alfalfa may have developed several strategies against Fe-deficient stress including the exudation of organic compounds like alfafuran which accelerate the Fe³⁺-reducing activity on the root cell membrane to dissolve insoluble iron compounds. Suzuki et al. (1995, 1997) observed that in barley several peptide spots obtained by electrophoresis were induced under Fe-deficient stress when mugineic acid-family phytosiderophores were secreted from the roots. They suggested that these peptides control the mugineic acid synthesis and secretion. We examined the peptides induced in Fe-deficient alfalfa roots.     

Tuber morphology,germination behaviour and propagation efficiency in three edible <Emphasis Type="Italic">Momordica</Emphasis> (Cucurbitaceae) species of India

Tuber morphology and propagation efficiency in three dioecious, tuber bearing Momordica species of India were investigated. Tuber morphology of M. dioica and M. sahyadrica is different from that of M. subangulata ssp. renigera. M. dioica and M. sahyadrica develop taproot tuber with shoot sprouts only at caudex region, whereas M. subangulata ssp. renigera has both taproot and adventitious tubers with shoot sprouts all over tuber surface. Tubers of M. dioica and M. sahyadrica are perennating whereas, in M. subangulata ssp. renigera they serve as both perennating and propagation structures. M. dioica and M. sahyadrica had prolonged tuber dormancy of 4–5 months whereas, M. subangulata ssp. renigera had short dormancy period of about 2 months. Cutting of tuberous roots is a highly efficient method for multiplication in M. subangulata spp. renigera, whereas whole tubers or longitudinal splits in to two or at best four equal halves with a portion of the apical meristem may work in the case of M. dioica and M. sahyadrica.     

Ecotoxicological evaluation of Harbour sediments using marine organisms from different trophic levels

Background, Aim and Scope  The toxicity of contaminated sediments should be evaluated considering the direct exposure of laboratory organisms to whole sediments and the indirect exposure to elutriates or extracts (Tay et al. 1992, Byrne and Halloran 1999, Nendza 2002). The alga Dunaliella tertiolecta is indicated for the use in toxicity bioassays because it is highly sensitive to several xenobiotics. Harpacticoid copepods have been already used for toxicity testing and Tigriopus fulvus is a promising Mediterranean target-species in ecotoxicology (Todaro et al. 2001, Faraponova et al. 2003, Pane et al. 2005a). In this study, the toxicity of sediments collected in harbour sites of the Northeastern Adriatic Sea was evaluated by growth inhibition test with free living and alginate-immobilized Dunaliella tertiolecta and acute toxicity test with nauplii and adult Tigriopus fulvus with the aim of pointing out the importance to utilize model organisms from different trophic levels in sediment ecotoxicology. Methodology  Elutriates and whole sediments were tested on free living and immobilized (Pane et al. 1998) algal cells, and on laboratory reared copepods. Free-living D. tertiolecta were exposed to diluted elutriates in a static, multi-well plate system. Naalginate immobilized D. tertiolecta were placed in polystyrene inserts fitted with polyester mesh bottoms and exposed to a thin layer (2 mm) of whole sediments in multi-well plates (EPS 1992, Pane and Bertino 1999). Toxicity tests with copepods were carried out on Tigriopus fulvus nauplii (elutriates) and adults (whole sediments and elutriates). Same-aged nauplii useful for toxicity tests were obtained by egg sac detaching and consequent hatching stimulation (Pane et al. 2006). Newborn nauplii (I–II stage) were exposed to elutriates in multi-well plates provided with polystyrene inserts. Adult T. fulvus maintained in polystyrene inserts fitted with polyester mesh bottoms were placed in contact with a thin layer (2 mm) of whole sediment placed on multi-well plate bottoms. All end-points were evaluated after 96 h. Results  In general, the effects increased with the increasing of elutriate concentration up to 50%; the stimulation or inhibition of algal growth was statistically significant in comparison to the control. The inhibiting elutriates induced EC₅₀ variations of algal growth ranging from 66.9% to 74.3%. The mortality of T. fulvus nauplii was always < 25% after treatment with 100% elutriates and < 10% after treatment with 50% dilution; no effect was shown up with 25% dilution; therefore LC₅₀ was not calculable. The effect of elutriates was negligible on adult copepods and LC₅₀ values were never calculable; percent mortality always resulted in < 10% after treatment with whole sediments. Discussion  Both experimental systems gave substantially similar results after exposition to whole sediments and elutriates. During the experiment with algal cells, the immobilization in Na-alginate and the employment of inserts which allowed the contact of organisms with sediments and their easy counting were particularly useful. Likewise, the employment of inserts of adequate mesh size in the tests with copepods allowed the contact of organisms with the sediment and made organism handling and counting easy, as well as the evaluation of mortality. The methodology here described and the utilization of the proposed test-species could have an importance also considering that the current trend in ecotoxicological research is towards finding the most appropriate organism for specific areas of concern by using indigenous species (Mariani et al. 2006) and towards the major significance of chronic and reproductive end-points. Conclusions  Based on the above results, it can be stated that the bioassay with Dunaliella tertiolecta could be a good estimation tool for the ecotoxicological assessment of marine sediments. The immobilization of algae in Na-alginate was seen to be useful to evaluate the toxicity of whole sediments; the employment of polystyrene inserts allowed an improvement of the procedures. T. fulvus nauplii and adults, as other harpacticoids such as Tigriopus japonicus (Yoon et al. 2006), satisfy the basic criteria for the employment of a standard species in marine bioassays. To date only pelagic Acartia tonsa are utilized in the standardized procedure to evaluate the risk assessment of chemicals or wastewaters (ISO 1999). As, on the contrary, the exposure of copepods to solid-phase contaminants it is not yet standardized, the employment of polystyrene inserts improved the procedures for T. fulvus too. So, the rapidity and the possibility to solve practical problems could be the main attractive features of this technique (Pane et al. 2005a) when applied to whole sediments. Recommendations and Perspectives  The methodology here developed being also applicable to long term and reproduction tests should be recommended because it provides relevant information in comparison with other frequently applied, standardized biotests with crustaceans (ISO 1999). The procedure has been shown to be easily applicable to selected marine organisms. ESS-Submission Editor: Prof. Dr. Henner Hollert (henner.hollert@bio5.rwth-aachen.de)     

Potential of Hydroxamic Acids in Breeding for Aphid Resistance in Wheat

Abstract

Swedish wheats were analysed at the seedling stage for hydroxamic acids (Hx), a family of natural aphid resistance factors. Analysis comprised the historical development from old landraces to modern cultivars, and included cultivars in the Swedish National List of Cultivars for 1992–93. Spring and winter wheats contained similar average Hx concentrations, 2.12 ± 0.801 (n = 26) and 2.37 ± 0.816 (n = 49) mmol/kg fr. wt, respectively. Cultivars recommended for 1992–93 showed lower levels of Hx, 1.14 ± 0.574 (n = 7) and 1.97 ± 0.751 (n = 8), respectively. Breeding wheat for higher Hx levels as a means of obtaining increased aphid resistance is discussed.     

Potential for leaching of potassium and phosphorus from pine and grass roots

Abstract

Nutrient analysis of roots recovered by wet sieving may be biased by leaching which occurs during sampling. The objective of this experiment was to determine the potential for leaching of potassium (K) and phosphorus (P) from roots of slash pine (Pinus elliottii Engelm. var elliottii) and Panicum aciculare grass seedlings. Roots of both species were sampled by wet and dry methods. The wet method included soaking for 15 d in deionizd water. All samples were dried prior to measuring their contents of sand, K and P. The amount of sand retained by both types of roots was reduced by wet treatment. Concentrations of K and P in both species also were significantly lower for wet‐treated roots than for dry roots. Multipliers for converting concentrations in wet‐treated roots to concentrations in dry roots were, for pine; 5.4 and 1.2 for K and P, respectively, and for grass 5.9 and 3.1 for K and P, respectively. However, these factors may be inappropriate for roots that had less exposure to water, or to roots of different anatomy and nutrient concentrations.     

Effect of Root Zone Temperature on Accumulation of Molybdenum and Nitrogen Metabolism in Potato Plants

Abstract

Changes in root temperature caused by the application of plastic covers were studied in relation to the uptake and content of molybdenum (Mo) in the different organs of potato (Solanum tuberosum L. var. Spunta) plants (roots, tubers, stems, and leaves) and in relation to nitrogen (N) metabolism. For the semi‐forcing technique of mulching, four different covers were used: T ₁ (transparent polyethylene), T ₂ (white polyethylene), T ₃ (coextruded black and white polyethylene), and T ₄ (black polythylene). The control treatment had no mulch. The results revealed a positive and significant effect of plastic covers on root temperatures: T ₀ = 16°C, T ₁ = 20°C, T ₂ = 24°C, T ₃ = 27°C, T ₄ = 30°C. These thermal differences significantly influenced the Mo concentration, particularly in the T ₂ and T ₃ treatments in the leaves, roots, and tubers. The same temperatures significantly altered N metabolism in both the aerial and underground parts of the plants, and a strong interrelationship was found between Mo and nitrate reductase (NR) activity. The mulching of this crop proved to be a promising technique in phytoremediation.     

Effect of placement of coated urea fertilizer on root growth and rubidium uptake activity in soybean plant

Abstract

We reported in the previous paper (Takahashi et al. 1991) that the deep placement of slow release N fertilizer (coated urea) contributed to a stable increase of soybean (Glycine max L. Merr.) yield. In the previous study we observed that the deep placement of coated urea did not depress appreciably the nitrogen fixation by root nodules although fertilizer N was efficiently utilized. We assumed that the N absorbed from the roots in the deep layers did not cause nodule senescence, contributed to the maintenance of the leaf activity during the maturation stage, and that the increase in the availability of carbohydrate and N improved seed production. In the current report the effects of placement of coated urea fertilizer on the root growth and activity were studied by measuring the root dry weight and Rb absorption activity.     

Stoichiometry for the decrease in the amount of phosphate-induced negative charges of clay fractions of andisols

Abstract

It is well-known that the amount of negative charges increases with phosphate (P) sorption on soils or soil constituents (Mekaru and Uehara 1972; Schalscha et al. 1972). The increase in the amount of negative charges could be considered as one of the beneficial aspects of P dressing for the retention of cationic nutrients.     

Quantification of the length and diameter of root segments with public domain software

Abstract

Roots control the uptake of water and nutrients by plants, and hence their dry matter production. One of the greatest constraints to our understanding of root dynamics has been the laborious tedium associated with accurate measurements of various root parameters. A technique is presented for the fast and accurate measurement of the length and diameter of all fragments of corn (Zea mays L.) roots contained in most 0.04 by 0.15 m soil cores by analyzing a single NIH‐Image of those roots. These parameters are determined by measuring the perimeter and area of each root fragment utilizing NIH‐Image analyses contained in a public domain software (NIH‐Image). When length and diameter are coupled with nutrient uptake data, we will have a better understanding of where plants extract nutrients and water in space and time.     

Effects of Cd on the fluxes of K+ and H+ in excised roots

As a result of Cd treatment, K concentrations decreased in Cd sensItive maize and kidney bean calli (Obata et al. 1994) and in intact roots of kidney bean plants (Obata et al. unpublished). Potassium may be extruded from the roots or the absorption of K may be depressed by the Cd treatment in these Cd sensitive plants. Obata et al. (1996) observed that Cd inhibited both the efflux of H⁺ and influx of K⁺ following K⁺ addition in intact roots of bean. Thus Cd may affect the activity of proteins essential to ion movement., i.e. ioncarriers, channels and ATPase embedded in the membranes and/or may affect the permeability of the lipids of the membrane.