枫香秋季变色期叶色变化及其生理基础

为探究秋季枫叶呈色的关键生理因素,该文以转色期叶色为绿色、黄色和红色的枫香单株为试材,研究了L^*、a^*、b^*值变化与叶片色素、可溶性糖及可溶性蛋白质含量变化的相关性。结果表明:(1)在变色期,3种色彩枫香叶片叶绿素a、叶绿素b、总叶绿素和类胡萝卜素均大量降解,花色素苷不同程度积累。(2)绿色叶单株叶绿素和类胡萝卜素始终保持较高含量,花色素苷含量上升4.2倍,叶片内色素含量比值始终保持稳定; 黄色叶单株叶绿素和类胡萝卜素含量最低,花色素苷含量上升4.4倍,b^*值与叶绿素含量极显著负相关,与类胡萝卜素含量显著负相关,与花色素苷/类胡萝卜素含量比值极显著正相关; 红色叶单株叶绿素和类胡萝卜素含量略高于黄色叶单株,花色素苷含量上升27.2倍,a^*值与叶绿素含量、类胡萝卜素含量极显著负相关,与花色素苷含量显著正相关,与色素含量比值无显著相关性。(3)红色叶单株具有较高的可溶性糖含量和可溶性蛋白质含量。因此,在枫香叶片变色期,保持较高的叶绿素和类胡萝卜素含量,维持色素含量比值稳定使叶片呈现绿色; 叶绿素和类胡萝卜素的大量降解,以及花色素苷/类胡萝卜素含量比值的升高使叶片呈现黄色; 叶绿素的降解和花色素苷的大量合成使叶片呈现红色。     

不同叶色红栌叶片中色素含量、酶活性及内含物差异的研究

为明确秋季不同叶色美国红栌叶片的生理差异,以秋季同一植株上红色、中间色、绿色三种颜色的美国红栌叶片为试材,测定了叶片中色素物质含量、酶活性以及叶片可溶性内含物的含量,结果表明：在红色叶片中,叶绿素含量较低,PAL、POD酶活性较高,花青素苷/叶绿素的比值较大,从而使叶色显现红色;而在绿色叶片中,叶绿素含量较高,PAL、POD酶活性较小,花青素苷/叶绿素的比值较低,叶片显现绿色。通过可溶性内含物测定可知,在红色叶片中的可溶性糖和蛋白质含量相对较高,均与花青素苷/叶绿素的比值达到显著相关水平,表明这些内含物的积累有利于花色素苷的合成。     

Protection mechanisms against excess light in the resurrection plants Craterostigma wilmsii and Xerophyta viscosa

Mechanisms of avoidance and protection against light damage were studied in the resurrection plants Craterostigma wilmsii and Xerophyta viscosa.In C. wilmsii, a combination of both physical and chemical changes appeared to afford protection against free radical damage. During dehydration leaves curled inwards, and the abaxial surface became exposed to light. The tissue became purple/brown in colour, this coinciding with a three-fold increase in anthocyanin content and a 30% decline in chlorophyll content. Thus light-chlorophyll interactions are progressively reduced as chlorophyll became masked by anthocyanins in abaxial layers and shaded in the adaxial layers. Ascorbate peroxidase (AP) activity increased during this process but declined when the leaf was desiccated (5% RWC). During rehydration leaves uncurled and the potential for normal light-chlorophyll interaction was possible before full hydration had occurred. Superoxide dismutase (SOD) and glutathione reductase (GR) activities increased markedly during this stage, possibly affording free radical protection until full hydration and metabolic recovery had occurred.In contrast, the leaves of X. viscosa did not curl, but light-chlorophyll interactions were minimised by the loss of chlorophyll and dismantling of thylakoid membranes. During dehydration, free radical protection was afforded by a four-fold increase in anthocyanin content and increased activities of AP, GR and SOD. These declined during rehydration. It is suggested that potential free radical damage may be avoided by the persistence of anthocyanins during the period of thylakoid membrane re-assembly and full chlorophyll restitution which only occurred once the leaves were fully rehydrated.     

Structural and functional characteristics of photosynthetic apparatus of chlorophyll-containing grape vine tissue

Photosynthesis in tissues under periderm of woody stems and shoots of perennial plants occurs in environment that is very different from the internal environment of leaf chloroplasts. These tissues are characterized by high CO₂ and low O₂ concentrations, more acidic surroundings, besides that only light which have passed through periderm reaches photosynthetic antennas. In contrast to leaves of deciduous plants chlorenchyma tissues of wintering plant organs are exposed to temperature fluctuations during all seasons, that is why the photosynthetic apparatus of woody stems has to be able to adapt to a wide range of environmental temperatures. In order to reveal unique features, which enable photosynthetic apparatus of chlorenchyma cells in woody plant organs to implement biological functions under different light and temperature conditions, we studied photosynthetic tissues of stem cortex in grapevine (Vitis vinifera L.) under normal conditions and after exposure to suboptimal temperatures and high light intensity. Comparative analysis of photosynthetic pigment composition and low-temperature chlorophyll fluorescence emission spectrum of leaves, young shoots and chlorenchyma of lignified shoots revealed relatively high level of chlorophyll b and carotenoids, and high photosystem II (PSII) to photosystem I (PSI) ratio in woody shoots. Analysis of parameters of variable chlorophyll fluorescence revealed high PSII activity in grapevine shoot cortex and demonstrated improved freeze tolerance and higher sensitivity to light of photosynthetic apparatus in grape vine in comparison to leaves. It was shown for the first time that photosynthetic apparatus in chlorenchyma cells of vine undergoes so-called “state-transition”–fast rearrangements leading to redistribution of energy between photosystems. Analysis of fatty acid (FA) compositions of lipids in examined tissues showed that the FA unsaturation index in green tissue of vine is lower than in leaves. A distinct feature of FA compositions of lipids in vine cortex was relatively high level of linoleic acid.     

Coordination of anthocyanin decline and photosynthetic maturation in juvenile leaves of three deciduous tree species

Juvenile leaves in high-light environments commonly appear red as a result of anthocyanin pigments, which play a photoprotective role during light-sensitive ontogenetic stages. The loss of anthocyanin during leaf development presumably corresponds to a decreased need for photoprotection, as photosynthetic maturation allows leaves to utilize higher light intensities. However, the relationship between photosynthetic development and anthocyanin decline has yet to be quantitatively described. In this study, anthocyanin concentration was measured against photopigment content, lamina thickness, anatomical development, and photosynthetic CO(2) exchange in developing leaves of three deciduous tree species. In all species, anthocyanin disappearance corresponded with development of c. 50% mature photopigment concentrations, c. 80% lamina thickness, and differentiation of the mesophyll into palisade and spongy layers. Photosynthetic gas exchange correlated positively with leaf thickness and chlorophyll content, and negatively with anthocyanin concentration. Species with more rapid photosynthetic maturation lost anthocyanin earliest in development. Chlorophyll a/b ratios increased with leaf age, and were lower than those of acyanic species, consistent with a shading effect of anthocyanin. These results suggest that anthocyanin reassimilation is linked closely with chloroplast and whole-leaf developmental processes, supporting the idea that anthocyanins protect tissues until light processing and carbon fixation have matured to balance energy capture with utilization.     

Uptake and release of nutrients by living and decomposing Typha glauca godr. tissues at Eagle Lake,Iowa

Uptake and release of nitrogen, phosphorus, potassium, calcium, and sodium from living above-ground and below-ground tissues and from decomposing litter of Typha glauca Godr. were studied at Eagle Lake, IA, during 1976. All nutrients were accumulated rapidly by shoots in the spring. Some of the nitrogen and phosphorus came from belowground storage; but potassium, calcium, and sodium were extracted entirely from the soil. Nutrients were immobilized in shoot tissues for different periods of time. Potassium content declined as rapidly as it had accumulated, and there was no evidence of belowground storage. Nitrogen and phosphorus content also declined, though not as rapidly. Approximately 45% of the nitrogen and phosphorus lost from the shoots was translocated to the rhizomes and stored. Calcium and sodium were conserved in shoot tissues until the shoots died. In the decomposing litter, potassium and sodium content declined, phosphorus and calcium content remained relatively constant, but nitrogen content increased. Over the full year of production and decomposition, this Typha glauca stand accumulated calcium and nitrogen, maintained phosphorus levels, and lost potassium and sodium.     

A system in which anthocyanin synthesis is induced in regenerated torenia shoots

A system in which anthocyanin synthesis can be induced under defined conditions was established in regenerated torenia shoots. Leaf discs prepared from torenia plantlets grown under sterile conditions were placed on solidified half-strength MS medium containing 3% sucrose and 4.4×10^–6 M benzyladenine (BA) and cultured under 16 h light/8 h dark (standard light) conditions for 10 days, then in the dark for a further 10 days. The discs were transferred to medium containing 7% sucrose without BA and cultured under standard light conditions. Six days after transfer, anthocyanin synthesis started in the regenerated shoots, and thereafter, anthocyanin accumulation increased while chlorophyll content decreased. Experiments in which either the timing of illumination was altered or shoots were retransferred to medium containing 1.5% sucrose or other sugars as well as sucrose indicated that both osmotic stress and light are required to induce anthocyanin synthesis. Once anthocyanin synthesis was induced in the torenia shoots 6 days after transfer, the shoots were fated to the synthesis of anthocyanins and the degradation of chlorophylls, and could not revert to the developmental pathway of shoot regeneration. This system may provide a good model for the investigation of the mechanisms underlying the induction of anthocyanin synthesis.     

Effect of ABA upon anthocyanin synthesis in regenerated torenia shoots

To elucidate the mechanism of anthocyanin synthesis induction concomitant with chlorophyll degradation, we established a system in which anthocyanin synthesis and degradation of chlorophyll in regenerated torenia (Torenia fournieri) shoots was induced on medium containing 7% sucrose. Here, we studied the effect of several plant-growth regulators on anthocyanin synthesis and the degradation of chlorophyll in the torenia shoot regenerating system. Exogenous abscisic acid (ABA) could induce anthocyanin synthesis and chlorophyll senescence in regenerating torenia shoots on the medium containing a low concentration of sucrose (1.5%). We determined the changes in the amount of endogenous ABA in the regenerated shoots during the process of anthocyanin synthesis on the medium containing 7% sucrose. It was revealed that the 7% sucrose treatment elevated endogenous ABA levels before the induction of anthocyanin synthesis and chlorophyll degradation. However, while retransfer to the 1.5% sucrose medium resulted in a gradual decrease in the ABA level and a failure of induction of anthocyanin synthesis, normal shoot regeneration. These results suggest that changes in the amount of endogenous ABA may play an important role in the induction of anthocyanin synthesis and chlorophyll degradation in regenerated torenia shoots.     

Regulatory effect of stems on sucrose-induced chlorophyll degradation and anthocyanin synthesis in Egeria densa leaves

Detached green leaves of the aquatic plant Egeria densa showed chlorophyll degradation and turned red due to induced anthocyanin synthesis incubated in 0.1 M sucrose under continuous light for 7–10 days. If the leaves were placed in water, only chlorophyll degradation occurred and the detached leaves turned yellow. The levels of endogenous total carbohydrates increased in detached leaves cultured in the sucrose solution but only increased marginally in water. If the leaves were still attached to a piece of stem, with a node on either side of the single leaf whorl, then they did not accumulate anthocyanin in culture with 0.1 M sucrose. These leaves showed a similar increase in total carbohydrates and degradation of chlorophyll as detached leaves. Attached leaves, in which the midrib had been cut in situ, showed localized accumulation of anthocyanin in the leaf tissue distal to the cut in the midrib when cultured in 0.1 M sucrose. These results suggest that the stem plays a regulatory role in anthocyanin synthesis in attached leaves cultured in a sucrose solution but does not influence chlorophyll degradation or carbohydrate accumulation.     

Morphology and photosynthetic physiology of Grimmia antarctici from wet and dry habitats

Summary Photosynthetic capacity, chlorophyll content and leaf and cell morphology were compared in Grimmia antarctici from wet and dry sites in the Bailey Peninsula SSSI, near Casey Station, East Antarctica. In wet sites G. antarctici grew as a turf with tall shoots of loosely packed long leaves: in very dry sites it formed small cushions with short shoots of small tightly packed leaves. Intermediate forms (large cushions) were also frequently observed in less extreme situations. Cell size and number were greater in drier sites. The chlorophyll content, chlorophyll a/b ratio and the light saturated photosynthetic and dark respiration rates at full turgor and under enhanced conditions of CO₂ were the same. This rules out a direct effect of water stress on the integrity of the photosynthetic apparatus and implies that the cushion form is a product of direct effects of water availability on cell division and differentiation and CO₂ assimilation under field conditions.     

Nondestructive estimation of anthocyanins and chlorophylls in anthocyanic leaves

The anthocyanin and chlorophyll contents in leaves provide valuable information about the physiological status of plants. Thus, there is a need for accurate, efficient, and practical methodologies to estimate these biochemical parameters of vegetation. In this study, we tested the performance and accuracy of several nondestructive, reflectance-based techniques for estimating anthocyanin and chlorophyll contents in leaves of four unrelated species, European hazel (Corylus avellana), Siberian dogwood (Cornus alba =Swida alba), Norway maple (Acer platanoides), and Virginia creeper (Parthenocissus quinquefolia), with widely variable pigment content and composition. An anthocyanin reflectance index, which uses reflectances in the green and red edge spectral bands, and a modified anthocyanin reflectance index, employing, in addition, the near-infrared (NIR) band, were able to accurately estimate leaf anthocyanin for all species taken together with no reparameterization of algorithms. Total chlorophyll content was accurately estimated by a red edge chlorophyll index that uses spectral bands in the red edge and the NIR. These approaches can be used to estimate anthocyanin and chlorophyll nondestructively and allow the development of simple handheld field instrumentation.     

Pigment variation and transcriptional response of the pigment synthesis pathway in the S2309 triple-color ornamental kale (Brassica oleracea L. var. acephala) line

Ornamental kale is popular because of its colorful leaves and few studies have investigated the mechanism of color changes. In this study, an ornamental kale line (S2309) with three leaf colors was developed. Analysis of the anthocyanin, chlorophyll, and carotenoid contents and RNA-seq were performed on the three leaf color types. There was less chlorophyll in the white leaves and purple leaves than in the green leaves, and the anthocyanin content was greatest in the purple leaves. All the downregulated DEGs related to chlorophyll metabolism were detected only in the S2309_G vs. S2309_W comparison, which indicated that the decrease in chlorophyll content was caused mainly by the inhibition of chlorophyll biosynthesis during the leaf color change from green to white. Moreover, the expression of 19 DEGs involved in the anthocyanin biosynthesis pathway was upregulated. These results provide new insight into the mechanisms underlying the three-color formation.     

Identification and characterization of genes induced for anthocyanin synthesis and chlorophyll degradation in regenerated torenia shoots using suppression subtractive hybridization, cDNA microarrays, and RNAi techniques

Anthocyanin synthesis and chlorophyll degradation in regenerated torenia (Torenia fournieri Linden ex Fourn.) shoots induced by osmotic stress with 7% sucrose were examined to identify the genes regulating the underlying molecular mechanism. To achieve this, suppression subtractive hybridization was performed to enrich the cDNAs of genes induced in anthocyanin-synthesizing and chlorophyll-degrading regenerated shoots. The nucleotide sequences of 1,388 random cDNAs were determined, and these were used in the preparation of cDNA microarrays for high-throughput screening. From 1,056 cDNAs analyzed in the microarrays, 116 nonredundant genes were identified, which were up regulated by 7% sucrose to induce anthocyanin synthesis and chlorophyll degradation in regenerated shoots. Of these, eight genes were selected and RNAi transformants prepared, six of which exhibited anthocyanin synthesis inhibition and/or chlorophyll degradation in their leaf discs. Notably, the RNAi transformants of the glucose 6-phosphate/phosphate translocator gene displayed inhibition both of anthocyanin synthesis and chlorophyll degradation in both leaf discs and regenerated shoots. There was also less accumulation of anthocyanin in the petals, and flowering time was shortened. The genes we identified as being up-regulated in the regenerated torenia shoots may help further elucidate the molecular mechanism underlying the induction of anthocyanin synthesis and chlorophyll degradation.     

氮素形态和铁营养对玉米苗期生长及体内铁分布的影响

以玉米(Zea mays)品种‘豫玉-22’为材料,采用营养液培养方法,研究了低铁和正常供铁条件下供应不同形态氮素对玉米苗期生长及体内铁分布的影响。结果表明:(1)与低铁介质相比,常铁介质增加了各氮素处理玉米幼苗的株高、地上部干重、全株干重,降低了根冠比,其中硝态氮处理表现得尤其突出;与供应硝态氮(NO3--N)相比,增施铵态氮(1/2 NO3--N 1/2 NH4 -N,NH4 -N)能明显促进低铁介质中玉米生长,但在常铁介质下作用不明显。(2)相比于低铁介质,正常供铁显著提高了相应处理玉米新叶叶绿素含量及净光合速率;2种供铁介质中,NH4 -N处理的新叶叶绿素含量以及净光合速率均高于其它氮素处理。(3)相比于低铁介质,正常供铁处理总体上增加了玉米各部分活性铁含量和全铁含量,对NO3--N处理的新叶活性铁含量增加尤其明显;2种供铁介质中,NH4 -N均有利于提高新叶活性铁含量和植株地上部全铁含量。(4)玉米新叶活性铁含量不仅与其叶绿素含量显著正相关(r=0.979**),也与叶片净光合速率显著正相关(r=0.950**)。研究发现,供铁状况显著影响玉米新叶的叶绿素含量及其净光合速率且与供氮形态存在互作;供应铵态氮有利于提高缺铁条件下玉米新叶活性铁含量,增强玉米植株的光合能力,进而促进其正常生长。     

除内袋时间及摘叶对红富士苹果果实品质的影响

研究了红富士苹果除内袋时间、摘叶时间和摘叶数量对冠层光照条件、果实品质及一年生枝条贮藏营养的影响.结果表明:除内袋时间适当提前可明显提高单果质量、果皮花青苷含量和果实着色面积(P<0.05),与9月30日(果实采前18 d)除内袋处理相比,9月24日(果实采前24 d)除内袋处理的单果质量、果皮着色面积和花青苷含量分别提高了10.98%、28.15%和13.44%,但除内袋时间对果实硬度、可溶性固形物和可滴定酸含量的影响差异不显著.当分别摘除果实周围15、30和45 cm以内的叶片后,果实周围的光照环境得到了明显的改善,3种摘叶处理均极显著提高了果实周围的相对光照强度(P<0.01),比摘叶前分别增加了70%、95%和115%;摘除果实周围30～45 cm以内的叶片有利于果皮着色和花青苷的积累;不同摘叶处理对枝条贮藏营养和第二年的萌芽率无显著影响.表明提前摘除内袋,并摘除离果实30～45 cm距离以内的叶片对果实外观及品质有明显促进作用.     

Influence of lignin on the methanization of lignocellulosic wastes

The effect on anaerobic digestion of reducing the lignin content of vine shoots to 1% (w/w), by treatment with sodium chlorite in an acid medium at 80°C, is reported. The yields of methane obtained were 240 ml of CH₄ g⁻¹ of VS (volatile solids) fed for untreated vine shoots, and 370 ml of CH₄ g⁻¹ of VS fed for treated vine shoots. A mathematical model was used to calculate the kinetic parameters H and μ, and the increased biodegradability of the substrate in which lignin had been removed was confirmed. A study of the mass balances of the process under optimum conditions (temperature = 35°C; loading rate of 1 g litre⁻¹ digester day⁻¹) enabled the percentage of degraded cellulose to be calculated (35·5% for untreated vine shoots, 81·5% for the treated vine shoots), as were the volumes of biogas and methane produced per gram of VS introduced (VS₁) and degraded. The blocking effect of lignin on the methanization process was confirmed.     

Senescence in Leaves of Acer pseudoplatanus L. and Parthenocissus tricuspidata Planch: II. Changes in Potassium and Sodium Content in Leaves and Leaf Discs of Acer

Changes in sodium and potassium content of the leaves of fiveAcer trees are recorded during two seasons. In general, sodiumcontents remained constant during leaf maturity and increasedduring senescence, whereas potassium contents increased duringleaf maturity and decreased during senescence. Short-term fluctuationsin sodium content during one season could be related to periodsof rainfall preceding some of the days of sampling. Experimentsin which excised leaves were washed in a spray of de-ionizedwater showed that the proportion of leaf potassium which couldbe removed increased as senescence progressed, and the quantityleached was approximately proportional to the area of the leafwhich had yellowed. With increasing leaf age, the amount of potassium which diffusedquickly out of discs cut from the leaves increased, whereasdiffusion of sodium became less. Discs cut from senescent leaveswere capable of absorbing sodium and it is suggested that thisability might account for the increase in sodium content ofAcer leaves during senescence.     

Spectral reflectance indices and pigment functions during leaf ontogenesis in six subtropical landscape plants

Pigment combinations are regulated during leaf ontogenesis. To better understand pigment function, alterations in chlorophyll, carotenoid and anthocyanin concentrations were investigated during different leaf development stages in six subtropical landscape plants, namely Ixora chinensis Lam, Camellia japonica Linn, Eugenia oleina Wight, Mangifera indica L., Osmanthus fragrans Lowr and Saraca dives Pierre. High concentrations of anthocyanin were associated with reduced chlorophyll in juvenile leaves. As leaves developed, the photosynthetic pigments (chlorophyll and carotenoid) of all six species increased while anthocyanin concentration declined. Chlorophyll fluorescence imaging of Φ_PSII (effective quantum yield of PSII) and of NPQ (non-photochemical fluorescence quenching) and determination of electron transport rate-rapid light curve (RLC) showed that maximum ETR (leaf electron transport rate), Φ_PSII and the saturation point in RLC increased during leaf development but declined as they aged. Juvenile leaves displayed higher values of NPQ and Car/Chl ratios than leaves at other developmental stages. Leaf reflectance spectra (400–800 nm) were measured to provide an in vivo non-destructive assessment of pigments in leaves during ontogenesis. Four reflectance indices, related to pigment characters, were compared with data obtained quantitatively from biochemical analysis. The results showed that the ARI (anthocyanin reflectance index) was linearly correlated to anthocyanin concentration in juvenile leaves, while a positive correlation of Chl NDI (chlorophyll normalized difference vegetation index) to chlorophyll a concentration was species dependent. Photosynthetic reflectance index was not closely related to Car/Chl ratio, while a structural-independent pigment index was not greatly altered by leaf development or species. Accordingly, it is suggested that the high concentration of anthocyanin, higher NPQ and Car/Chl ratio in juvenile leaves are important functional responses to cope with high radiation when the photosynthetic apparatus is not fully developed. Another two leaf reflectance indices, ARI and Chl NDI, are valuable for in vivo pigment evaluation during leaf development.     

In vitro Growth and Shoot Multiplication of Achras zapota in a Controlled Carbon Dioxide Environment

The culture vessels with multiplying shoots of Achras zapota L. on Schenk and Hildebrandt (SH) medium containing 8.88 M 6-benzylaminopurine (BAP) with or without sucrose were kept under varied CO₂ concentrations ranging from 0.6 to 40.0 g m^–3 using different concentrations of sodium bicarbonate (NaHCO₃), sodium carbonate (Na₂CO₃), potassium bicarbonate (KHCO₃), and potassium carbonate (K₂CO₃) in small acrylic chambers. Complete absence of carbon source caused death of shoots within 20 d. Under elevated concentrations of CO₂ (10.0 and 40.0 g m^–3) the shoots grew photoautotrophically on sucrose-free medium. The growth of cultures was better at 40.0 g (CO₂) m^–3 than on 3.0 % sucrose under ambient air of growth room. However, the best response was obtained at 10.0 g (CO₂) m^–3 and 3.0 % sucrose where maximum number of shoots, shoot length, fresh and dry mass, total number of leaves and leaf area was observed.     

Sodium Absorption and Distribution in Forage Grasses of Different Potassium Status

The absorption and distribution of sodium were examined in threegrasses grown in flowing solution culture with different suppliesof potassium. There were marked differences between the speciesin the rate of absorption by their roots, timothy absorbingat a much slower rate than either ryegrass or cocksfoot. Inall species, the rate of Na absorption was greatest when therewas a maintained supply of K and/or when the K contents of theplants were high. Transport of Na from roots to shoots of timothywas restricted; it was less restricted in the other speciesand large proportions of the Na moved from roots to shoots whenK was not supplied to the plants. Sodium transported from theroots accumulated in old leaves and not in the younger leaves.When K was no longer supplied, the growth of ryegrass was maintainedin the plants previously grown with Na plus K; Na supplied insteadof K, however, did not maintain growth. Cocksfoot grown withNa grew less well than when grown without Na when plants wereno longer supplied with K; the growth of timothy was unaffectedby Na. Dactylis glomerata L., Lolium perenne L., Phleum pratense L., cocksfoot, ryegrass, timothy, absorption of ions, distribution of ions, potassium, sodium