Phosphorus enhances Al resistance in Al-resistant Lespedeza bicolor but not in Al-sensitive L. cuneata under relatively high Al stress

Background and Aims

Aluminium (Al) toxicity and phosphorus (P) deficiency often co-exist in acidic soils and limit crop production worldwide. Lespedeza bicolor is a leguminous forage species that grows very well in infertile, acidic soils. The objective of this study was to investigate the effects of Al and P interactions on growth of Lespedeza and the distributions of Al and P in two different Al-resistant species, and to explore whether P can ameliorate the toxic effect of Al in the two species.

Methods

Two species, Lespedeza bicolor and L. cuneata, were grown for 30 d with alternate Al and P treatments in a hydroponics system. Harvested roots were examined using a root-system scanner, and the contents of Al, P and other nutrient elements in the plants were determined using inductively coupled plasma-atomic emission spectroscopy (ICP-AES). Haematoxylin staining was used to observe the distribution of Al in the roots of seedlings. After pre-culture with or without P application, organic acids in the exudates of roots exposed to Al were held in an anion-exchange resin, eluted with 2 m HCl and then analysed using high-performance liquid chromatography (HPLC).

Key Results

Lespedeza bicolor exhibited a stronger Al resistance than did L. cuneata; Al exclusion mechanisms may mainly be responsible for resistance. P application alleviated the toxic effect of Al on root growth in L. bicolor, while no obvious effects were observed in L. cuneata. Much less Al was accumulated in roots of L. bicolor than in L. cuneata after P application, and the P contents in both roots and shoots increased much more for L. bicolor than for L. cuneata. Lespedeza bicolor showed a higher P/Al ratio in roots and shoots than did L. cuneata. P application decreased the Al accumulation in root tips of L. bicolor but not in L. cuneata. The amount of Al-induced organic acid (citrate and malate) exudation from roots pre-cultured with P was much less than from roots without P application; no malate and citrate exudation was detected in L. cuneata.

Conclusions

P enhanced Al resistance in the Al-resistant L. bicolor species but not in the Al-sensitive L. cuneata under relatively high Al stress, although P in L. cuneata might also possess an alleviative potential. Enhancement of Al resistance by P in the resistant species might be associated with its more efficient P accumulation and translocation to shoots and greater Al exclusion from root tips after P application, but not with an increased exudation of organic acids from roots.Key words: Lespedeza bicolor, L. cuneata, Al toxicity, Al resistance, root morphology, phosphorus     

ASYNCHRONOUS REPRODUCTIVE ACTIVITY IN THE BROADCAST SPAWNER CELLANA CAPENSIS (GMELIN, 1791) (GASTROPODA: PATELLIDAE)

Short-term fluctuations in the reproductive condition of thelimpet Cellana capensis are described. Gravimetric and histologicalanalyses of gonadal development were carried out on samplescollected at 4 day intervals over a 3 month period from theMkambati Nature Reserve, Transkei. The results indicate a distinctlack of reproductive synchrony within the population, whichapplied not only to gametogenic development per se but alsoto spawning activity. Some of the spawning events identifiedduring the study period were more marked than others, thesemay reflect a greater degree of synchronization in activity.Results presented suggest that C. capensis is probably a ‘partial’rather than a ‘complete’ spawner. The implicationsof asynchronous spawning activity for the reproductive successof broadcast spawners are also discussed. (Received 6 December 1988; accepted 15 February 1989)     

THE REPRODUCTIVE CYCLES OF THREE TROCHID GASTROPODS FROM THE TRANSKEI COAST, SOUTHERN AFRICA

Changes in gonadal histology were used to determine the reproductivecycles of three trochid gastropods commonly found along theTranskei coast. They were dioecious, the sexes being evenlydistributed both in terms of numbers and size. Despite someasynchrony, Monodonta australis had a distinct reproductivecycle with a short inactive period during the winter, gonadaldevelopment coincided with increasing sea temperatures and spawningtook place from late summer to autumn. Both Oxystele tabularisand O. variegata exhibited asynchronous intermittent spawningthroughout the study period. However, intensified spawning activityby O. tabularis was noted between April and September. The reproductivecycles and mechanisms of these trochids are compared with thoseelsewhere. (Received 20 January 1986;     

Secretion of malate and citrate from roots is related to high Al-resistance in Lespedeza bicolor

Lespedeza bicolor (Lespedeza bicolor Turcz. cv. Jiangxi) is a leguminous shrub that is well adapted to acid infertile soils. However, the mechanisms of aluminum resistance in this species have not been established. This study aimed to assess the possible resistance mechanisms of this plant to Al. An Al-sensitive species of Lespedeza, sericea lespedeza [Lespedeza cuneata (Dum.-Cours.) G. Don cv. Zhejiang], was used as a reference. The roots of L. bicolor secreted both malate and citrate after exposure to Al, but roots of L. cuneata did not. The secretion of organic acids from L. bicolor was specific to Al; neither 15-day P starvation nor 50 μM lanthanum induced the secretion of these organic acid anions. Secretion of organic acid anions in L. bicolor was detected after 3–6 h exposure to Al, and the amount increased significantly after 6 h exposure, suggesting that this plant shows a pattern II-type organic acid secretion. This is supported by the finding that the secretion was significantly inhibited by a protein-synthesis inhibitor, cycloheximide. Two kinds of anion-channel inhibitors had different effects on Al-induced secretion of organic acids: 9-anthracene carboxylic acid completely inhibited secretion, phenylglyoxal had no effect. Root elongation in L. bicolor was more severely inhibited by Al in the presence of 9-anthracene carboxylic acid. All these results indicated that the secretion of malate and citrate is a specialized response to Al stress in L. bicolor roots, which might be one of the Al-resistance mechanisms in this species.     

REPRODUCTIVE CYCLES OF CERASTODERMA GLAUCUM(BRUGUIERE) AND C. EDULE (L.) WITH SPECIAL REFERENCE TO THE EFFECTS OF THE 1981-82 SEVERE WINTER

Gonadal changes in two species of Cerastoderma occurring indiscrete populations in South Wales were monitored by stereologicalanalysis during the breeding seasons of 1981 and 1982. Fecundityand spawning efficiencies were determined quantitatively. Therelationship between reproductive activity and changes in meatcondition are discussed. Both species are opportunistic breeders. Their normal patternsof reproduction are similar, consisting of minimum activityin winter and peak activity during spring/early summer. Thetwo species were affected differently by the severe winter of1981–82—C. glau-cum exhibited a single, ‘epidemic’spawning and resumed gametogenesis after a 4-month resting period.C. edule adopted a ‘polycyclic’ pattern withouta resting period. Both of these unusual reproductive strategies,augmented by reduced predation resulted in heavy spatfalls. (Received 20 March 1985; revised 20 March 1985;     

滨海盐渍土壤中不同类型盐生植物富集镉的效应

为了利用被镉污染的滨海盐渍土壤,通过实验对比分析3种不同类型盐生植物对盐渍土中镉的富集效应,以期初步探明不同类型盐生植物在镉污染盐渍土壤修复中的效果。选择的3种盐生植物类型是:聚盐盐生植物,泌盐盐生植物和避盐盐生植物。通过温室盆栽实验,将植物在不同镉含量的盐渍土壤中种植培养60 d,测定和分析不同类型盐生植物对镉的生物浓缩因子、转移系数以及植株内地上部分和根部生物量和镉含量的变化。结果表明,不同镉含量的土壤对碱蓬和芦苇的生长影响较小,对二色补血草的生长影响较大。不同镉含量的土壤中,芦苇地上部分镉的生物浓缩因子变化差异不显著,并且其地上部分吸收镉的百分率较高。碱蓬和芦苇的转移系数大于二色补血草的转移系数,并且碱蓬的转移系数在不同镉含量的土壤中变化不显著;二色补血草的转移系数随着土壤中镉含量的增加而显著增大。3种盐生植物中,碱蓬最具修复镉污染盐渍土壤的潜力,这可能和它是聚盐盐生植物的生理类型有关。芦苇整个植株的地上部分富集镉的总量在3种植物中是最高的,因此,芦苇在镉含量较低时也可以做为镉污染盐渍土壤的修复材料。     

Nitrate Reductase Activity: A Biochemical Criterion of Hybrid Vigour in Sorghum bicolor (L.) Moench

F₁ hybrids of Sorghum bicolor (L.) Moench and their inbred parentswere analysed for NADH-nitrate reductase activity during theearly stages of seedling growth. In all the hybrids both mid-parentaland better parental heterosis were discernible in shoots whereasin roots two hybrids out of the three tested, showed heteroticlevels. It is suggested that in sorghum nitrate reductase activityduring seedling stages can be used as a biochemical criterionfor evaluating hybrid vigour. Sorghum bicolor (L.) Moench, sorghum, hybrid vigour, nitrate reductase     

扎龙湿地保护区异质生境芦苇种群分株构件的数量特征

采用单位面积取样,计数和测量的调查与统计方法,对扎龙湿地保护区4个生境单优群落芦苇种群分株构件数量特征进行比较分析。结果表明,4个生境芦苇种群从5月10日左右返青后进入营养生长期,分株高度和分株密度均持续增加到生殖生长初期的8月份,其中6-8月份差异均达到显著水平(P0.05),8-10月份差异均未达到显著水平(P0.05),芦苇进入生殖生长期后,分株便停止高度生长,地下芽的输出也不再形成分株补充现实种群;分株生物量和种群生物量均持续增加到生殖生长旺盛期的9月份,至休眠期的10月份均有所降低,各月份间的差异均达到显著水平,芦苇种群在生长季末期,具有将生产的物质分配转移到地下储藏与营养繁殖器官的形成与生长上的特性。芦苇种群分株数量特征与返青后实际生长时间之间均较好地符合对数函数关系,R~2在0.818-0.994之间,拟合方程均达到了P0.05的显著水平。4个生境芦苇种群分株密度以湿生生境最大,依次为水生生境、旱生生境,盐碱生境最小,分株高度、分株生物量和种群生物量均以水生生境最大,依次为湿生生境、旱生生境,盐碱生境最小。因此,4个生境芦苇种群分株构件数量特征均表现出基本一致的生育期节律性,同时,芦苇种群的个体生长和种群动态存在明显的环境效应,土壤含水量、pH是影响该地区芦苇分株数量特征的主要因子。     

Biochemical Basis of Heterosis in Sorghum: Changes in Chlorophylls and Ascorbic Acid Turnover during Seedling Growth

Shoots of three F1 hybrids of Sorghum bicolor (L.) Moench withtheir inbred parents were analyzed for chlorophyll a and chlorophyllb contents as well as for ascorbic acid turnover during theearly stages of seedling growth. In all the hybrids both chlorophylland ascorbic acid turnover showed better-parental and/or mid-parentalheterosis. Since ascorbate participates in photosynthesis andnitrate reduction, it is suggested that a well co-ordinatedsystem incorporating photosynthetic efficiency and nitrate assimilationmay be associated with the manifestation of hybrid vigour insorghum. Sorghum bicolor (L.) Moench, hybrid vigour, chlorophyll, ascorbic acid, nitrate reductases     

Interannual variability in reproductive traits of the Patagonian toothfish Dissostichus eleginoides around the sub‐Antarctic island of South Georgia

Commercial fisheries data, collected as part of an observer programme and covering the period 1997–2014, were utilized in order to define key reproductive traits and spawning dynamics of the Patagonian toothfish Dissostichus eleginoides at South Georgia. Multi‐year spawning site fidelity of D. eleginoides was revealed through the identification of previously unknown spawning hotspots. Timing of female spawning was shown to have shifted later, leading to a shorter spawning duration. A decrease in length and mass of female and male spawning fish and a reduced number of large spawning fish was found, evidence of a change in size structure of spawning D. eleginoides. During the study period fewer later maturity stage females (including spawning stage) were observed in conjunction with increased proportions of early stage female D. eleginoides. The findings are discussed in the context of reproductive success, with consideration of the possible effects such spawning characteristics and behaviours may have on egg and larval survival. This work presents the first long‐term assessment of D. eleginoides spawning dynamics at South Georgia and provides valuable knowledge for both the ecology of the species and for future fisheries management of this commercially important species.     

LC–MS-based chemotaxonomic classification of wild-type Lespedeza sp. and its correlation with genotype

In this study, 39 specimens belonging to Lespedeza species (Lespedeza cyrtobotrya, L. bicolor, L. maximowiczii, and Lespedeza cuneata) (Leguminosae) were classified phenotypically and genotypically. We constructed a phylogenetic tree based on the combined nrDNA (internal transcribed spacer; ITS) and cpDNA (trnL-trnF) sequences with the aim of classifying the genotypes. Samples were mainly divided into three genotypes. Samples of L. cyrtobotrya and L. bicolor were mixed in a single branch, whereas samples of L. maximowiczii and L. cuneata were clustered within species, respectively. We performed a liquid chromatography–electrospray ionization–mass spectrometry-based metabolite profiling analysis to classify the phenotypes. Multivariate statistical analyses such as principal component analysis (PCA) and hierarchical clustering analysis (HCA) were used for the clustering pattern analysis and distance analysis between species, respectively. According to the PCA and HCA results, leaves were classified into four phenotypes according to species. In both the genetic and chemotaxonomic classification methods, the distance between L. cyrtobotrya and L. bicolor was the closest between species, and L. cuneata was the farthest away from the other three species. Additionally, orthogonal partial least squares-discriminant analysis was employed to identify significantly different phytochemicals between species. We classified L. cyrtobotrya and L. bicolor by identifying significantly different phytochemicals. Interestingly, leaves and stems showed different phenotypic classifications based on the chemotaxonomic classification. Stem samples of the other three species were mixed regardless of species, whereas L. cyrtobotrya stem samples were clustered within species. The phenotypic classification of leaves coincided more with the genotypic classification than that of stems. Key message We classified four wild-type Lespedeza sp. by analyzing the combined nrDNA (ITS) and cpDNA (trnL-trnF) sequences. We also classified leaves and stems of Lespedeza sp. by applying liquid chromatography–mass spectroscopy-based metabolite profiling.     

PATTERNS OF REPRODUCTION IN FOUR LITTORINA SPECIES

The reproductive periodicities of four Iittorinids, Littorinaneglecta, L. nigrolineata, L. saxatilis and L. arcana were examinedin sympatric populations at Porth Swtan, Anglesey, North Wales.Three species, L. neglecta, L. nigrolineata and L. arcana, werefound to show marked reproductive periodicity. L. saxatilisshowed no periodicity and large numbers of animals were foundin breeding condition throughout the year. The oviparous speciesL. nigrolineata and L. arcana, came into reproductive conditionin the late summer or early autumn, L. nigrolineata in Augustand L. arcana in September, and their populations at Porth Swtanhad few individuals in breeding condition in the late springand early summer. L. neglecta, however, showed a periodicitywith high numbers of breeding animals throughout the springand summer and very low numbers in the autumn. For L. neglectathe basis of reproductive periodicity is shown to be the synchronizedannual life cycle of individuals of this species. It is thereforebelieved to differ from the other three species in which individualsare longer-lived and are supposed to undergo seasonal periodsof reproductive inactivity. (Received 2 March 1982;     

Description of Rhynchoteuthion larvae of Illex argentinus from the summer spawning subpopulation

The Rhynchoteuthion larvae of Illex argentinus resulting fromsummer spawning in North Patagonic shelf waters, its distributionand abundance, are described in this paper. The material wascollected in the Argentine Sea (35–55°S) by meansof plankton nets. The research cruises were made by the R/VShinkai Maru and the R/V Walther Herwig during the period April1978 to April 1979. The most important spawning ground of thesummer spawning subpopulation is found in continental-shelfwaters (between 43 and 46°S) during the period December-February.This area was established on the basis of both ripe (December)and spent females (February). The larvae which were caught duringthe same period, especially in March, confirmed the spawningarea of this demographic unit. The larvae showed the lengthof the mantle (ML) to be from 1.2 to 6.5 mm. Tentacles weresplitting in specimens from 5.0 to 6.5 mm ML (transition stage).When 7.0 mm ML or more, all specimens were juveniles and hadtheir tentacles completely separated. Larvae were characterizedas type C, following the proposal of Sato (1973) and Sato andSawada (1974) in the Bulletin of the Shizuoka Prefectural FisheriesExperimental Station.     

Maturation, Spawning and Development in Siphonaria Pectinata Linnaeus (Gastropoda: Pulmonata) at Gibraltar

A comparative study of some aspects of the reproductive biologyof Siphonaria pectinata was carried out at Gibraltar. Maturationoccurred at 5-6 mm shell length corresponding to an age of 6-12months. At the more sheltered Atlantic site spawning was evidentfrom March to June 1992 with peak spawning occurring in May-June1992. The patterns of spawning activity for the following twoyears were very similar. Egg ribbons were very scarce at themore exposed Mediterranean site and were found in considerablyless quantities than at the Atlantic site. Individuals at bothsites contained mature eggs in their gonads at the onset, andthen throughout the duration of the spawning season, but exhibitedspent gonads by October. The spawn of S. pectinata consistedof small oval ribbons filled with numerous egg-shaped capsulesspaced irregularly in a jelly-like matrix. Spawn was depositedmainly away from the home scar in shallow depressions, cracksand crevices and between barnacles and mussels. S. pectinataexhibited planktonic development. The fertilised eggs took between14 and 21 days to develop at ambient temperature (14-19°),and hatched as free-swimming planktotrophic veligers. The timingof spawning activity and the placement of egg masses in protectivemicrohabitats on the shore are discussed in terms of adaptationsto reduce desiccation stress, predation and possible food shortages.The mode of development in S. pectinata is placed in contextwith current theories regarding siphonariid ancestry and evolution. (Received 6 January 1998; accepted 25 June 1998)     

HETEROSTOPHIC SHELLS AND PELAGIC DEVOLOPMENT IN TROCHOIDEANS: IMPLICATIONS FOR CLASSIFICATION, PHYLOGENY AND PALAEOECOLOGY

Four species of archeogastropods, presumed members of threesubfamilies of the trochidae, exhibit significant differencesin developmental modes and shell coiling. All four species havelecithotrophic development which is reflected in their inflatedpausispiral protoconchis; however, Margarites marginatus andLirularia succincta have benthic development in gelatinous masses,while Margarites pupillus and Calliostoma ligatum have pelagicembryos and swimming larvae with a potential for dispersal overa period of a week or longer. These modes cannot be deducedfrom the size of the egg, the size or shape of the protoconch,or the size or relative prominence of female pallial reproductivestructures. The protoconch of C. ligatum is orthostrophicallycoiled, but the protoconchs of the other three species are hyperstrophicallycoiled although their teleconchs are orthostrophic. These threetrochoidean species thus share with architectonicoideans, pyramidelloideans,opisthobranchs, and pulmonates the distinctive shell characterof heterostrophy, previously unreported for archaeogastropods. These observations, considered together with others reportedin the literature, show: (1) that developmental mode (pelagicor benthic) cannot be inferred from protoconch appearance ortaxonomy in major trochoidean ganera; (2) that significant dispersalpotential is present in the histories of some trochoidean archeogastropoids;and (3) that character sets (pallial reproductive structures,pairing during spawning, heterostrophic shell coiling) thoughtnot to occur below the mesogastropod level are found in theTrochoidea. These conclusions bring into question the usefulnessof these characters in defining higher gastropod taxa and raiseadditional questions concerning the ancestry of the higher gastropods. (Received 16 February 1989; accepted 25 June 1989)     

Comparative Leaf Surface Morphology and the Glossy Characteristic of Sorghum, Maize, and Pearl Millet

Leaf surfaces of seven genotypes of Sorghum bicolor, two ofmaize, Zea mays, and two pearl millet, Pennisetum americanum,were examined by scanning electron microscopy for possible morphologicaldifferences. Leaves 1, 3, 5 and 7 were photographed and printswere used to estimate waxiness, hairiness or pubescence andstomatal density. Glossiness was determined by spraying water,which adhered to the glossy leaves. Cuticular transpirationof detached third and fifth leaves was estimated from the rateof water loss after abscisic acid induced stomatal closure.Sorghum lines SC283, CSM63, CSM90, and pearl millets Souna andTiotioni (all from Mali), were non-glossy, well covered withwax, and exhibited variable hairiness. Older leaves of sorghumvarieties Martin and Redlan were glossy and, like older leavesof the other glossy lines SC1096 and SC90, had little or nowax deposits on their cuticles. The two maize cultivars, NB611and N7A, were non-glossy with dense wax covering; no trichomeswere observed until the 5 to 7 leaf stage. Thus, the glossycharacter was correlated with the reduction or absence of waxdeposits on the leaf surfaces, while hairiness might occur ineither glossy or non-glossy genotypes. Unlike sorghum and maize,in which all leaves after the fifth or seventh were glossy,pearl millet showed no glossiness through the ninth leaf. Measurementsshowed that cuticular transpiration of glossy leaves was oftenmore than double that of non-glossy leaves. Comparisons amongsorghums showed that non-glossy lines had higher stomatal densitiesthan glossy lines. Epicuticular wax, trichome, glossy mutant, stomata, cuticular transpiration, Sorghum bicolor, (L.) Moench, Zea mays L., Pennisetum americanum, (L.) Leeke     

Temperature, food and mate limitation of copepod reproductive rates: separating the effects of multiple hypotheses

Temperature, food and the availability of mates may all limitrates of egg production is freshwater diaptomids. We have usedthe reproductive phases of Diaptomus paliidus to develop anindex of food limitation (f) and an index of mate limitation(M), each of which responds independently of temperature. Theresponse of the food limitation index to high and low temperaturesand high and low food concentrations was examined in a two-wayfactorial experiment in the laboratory. The index was highlyresponsive to a change in food concentration, stable duringa change in temperature alone at the high food level and responsiveto the synergistic interaction of food and temperature effectswhen both factors were limiting. Laboratory data indicate thatthe f index should not be biased by mate limitation unless maledensities are 6.7 x 10^–l0 males 1^–1 or less. Themate limitation index was highly responsive to mate availability,and ranged from a low of 9–19 in the presence of abundantmales to 100 in tbe absence of males. Applications of the fand M indices to a natural population of D. paliidus over 6weeks indicated that both food and mate availability were limitingthe reproductive rates of the copepods during the sampling period. ¹Present address: Institute of Animal Resource Ecology, Universityof British Columbia, Vancouver, Canada V6T 1W5     

Distribution and impact of exotic gall flies (Lipara sp.) on native and exotic Phragmites australis

Two exotic gall fly species infest stems of native and exotic Phragmites australis (Cav.) Trin. ex Steudel in northeastern North America. In this study, we determined the distribution of Lipara similis Schiner and L. rufitarsis Loew in native and exotic P. australis in Rhode Island. We also studied the within-stand distributions of each fly species and their effects on flowering of native and exotic P. australis. We collected stems from populations throughout southern Rhode Island and measured stem length and diameter, and percent flowering. Stems were then dissected to determine Lipara infestation. L. similis and L. rufitarsis were found throughout Rhode Island infesting both native and exotic P. australis, but their presence and abundance varied among sites. Within stands, L. similis infests the taller, thicker interior stems and L. rufitarsis infests the shorter, thinner exterior stems. Lipara similis reduces stem length by 6%; L. rufitarsis infestation reduces stem length by 37%. The flowering rate of uninfested stems is significantly lower in native P. australis stems than in exotic stems. Both Lipara species prevent infested stems from flowering. In adjacent stands of native and exotic P. australis, L. rufitarsis infests significantly more native stems than exotic stems, possibly further reducing the reproductive potential of the native plants relative to the exotic. Lipara species may play a role in facilitating the displacement of native P. australis by the exotic genotype.     

The Distribution of Silicon Deposits in the Roots of Molinia caerulea (L.) Moench. and Sorghum bicolor (L.) Moench.

The occurrence of silica in relation to meristematic zones andthe thickening of the endodermis in the roots of Molinia caerulea(L.) Moench. and Sorghum bicolor (L.) Moench. has been investigatedby means of the electron-probe microanalyser and the scanningelectron microscope. In proximal regions of mature roots ofM. caerulea, the central strengthening tissue of the stele,the vessel walls, the endodermis and the sub-epidermal sclerenchymaare areas of heavy accumulation. The distal regions of suchroots are relatively free of silicon and show little thickeningof the inner tangential walls of the endodermis or of the cellsof the strengthening tissues. The thickening of these elementsis shown to be associated with their location and the age ofthe root. In the proximal regions of S. bicolor, silicon is detected andlargely confined to the inner tangential walls of the endodermiswhich display some thickening. In addition, discrete and evenly-distributeddeposits varying in size partly fill the lumen of this layer.Some cells exhibit a number of smaller protrusions. High magnificationsof these lumen deposits show a distinct granular structure incontrast to the very uniform pattern of the wall deposits. The results are compared with deposits in grass leaves and inflorescencebracts and in woody perennials. The presence of silicon in additionto suberin, lignin and polyphenols in the thickened endodermalwall is also discussed in relation to the recognized functionof the endodermis.