text:
East Med Gas Forum wants worldwide upgrade
Energy ministers of the East Med Gas Forum (EMGF) met in Egypt on 25 July, agreeing to access the possible upgrade of the forum to the level of an international organisation, the first of its kind in southeast Mediterranean, and form a committee with the natural gas industry, which would include the participation of state and private companies from the forum member countries.
Egyptian Energy Minister Tariq el-Mulla and his counterparts Greece's Costis Hatzidakis, Cyprus' Yiorgos Lakkotrypis, Israel's Yuval Steinitz, Jordan's Hala Zawati, and Italy's Deputy Minister Andrea Cioffi attended the forum, which is supported by the United States and the European Union.
US Energy Secretary Rick Perry and Assistant Secretary of State for Energy Resources Frank Fannon attended the session on 25 July, which follows the joint declaration of the 20 March 2019 summit between Cyprus, Greece, and Israel, with the participation of the Secretary of State Mike Pompeo. At that session, Pompeo underlined Washington's support for the trilateral mechanism and noted the importance of increased cooperation.
Hydrocarbons Company CEO Charles Ellinas told New Europe on 29 July that the meeting formalised the setting up of the EMGF and agreed to set up committees to take its objectives forward. "However, it was useful in terms of regional politics but vague and lacking on practicalities, especially given the region's burning problems in need of urgent resolution," he said. "It is a talking shop and as such it will help improve cooperation among its members, but it will not have a direct effect on the development and export of hydrocarbons from the region," Ellinas said, adding that is still subject to the commercial challenges of global markets and the effects of the global shift to clean energy.
Greece, Cyprus and Israel to meet in Athens on 7 August
The European Commission announced in Egypt on 25 July that the EC would support the Forum's activities with significant financial assistance while the ministers also approved the first study to be conducted on behalf of EMGF in cooperation with the World Bank to examine the potential of the region's natural gas and possibilities for natural gas development and export, the Greek Energy Ministry said in a statement.
Hatzidakis also has a bilateral meeting with Perry and Fannon in Egypt where they discussed US-Greek energy relations and their geopolitical significance. The Greek Energy Minister highlighted the importance of the EastMed gas pipeline for Europe's energy security and called for the need to overcome the current delay due to the pending signing of the intergovernmental pipeline agreement between Greece, Cyprus, Israel and Italy, the Greek Energy Ministry said, adding that Fannon will lead the US delegation to the trilateral meeting of the energy ministers of Greece, Cyprus and Israel in Athens on 7 August. That meeting, according to Ellinas, is a continuation of previous tri-partite meetings between these countries with US support. "It will probably concentrate on the East Med gas pipeline, but the project can progress only if it can provide gas to Europe at prices that can compete with existing gas prices. This is where the challenge lies," he said.
Ellinas noted that Washington would like to see East Med gas exported to Europe to lessen dependence on Russian gas. But this can happen only if East Med gas can compete with prices prevalent in Europe, which is a challenge. However, LNG from Egypt's existing LNG plants at Idku and Damietta may be exported to Europe because very low liquefaction costs make it commercially viable, but other potential gas exports from the region cannot benefit from this advantage. This does not need the cooperation of Turkey," Ellinas said.
Turkey's role in the East Med
He noted that Turkey's impact is on what happens with exploration and export of gas from Cyprus' Exclusive Economic Zone (EEZ). "The EMGF cannot help with such problems, especially as Turkey is not a member of EMGF," Ellinas argued. "The EU and US might have been able to use their influence, but pressure and sanctions so far have not had any impact. It anything, it would appear that they have lessened their ability to influence Turkey, which says that it no longer sees the EU as an independent arbiter," he said.
At the Forum on 25 July in Egypt, Hatzidakis also discussed Greek-Egyptian energy cooperation with el-Mulla, focusing on marketing, exploration and production of hydrocarbons, regional cooperation in the gas sector as wells as investments in renewables and electricity.
El-Molla and Perry also met in Egypt with the latter reportedly stressing the role of Egypt as a reliable source of energy, pointing out the US readiness to provide all the expertise and technologies to Egypt to enhance its natural position as an important energy hub in the region, and highlighting Egypt's role as a cornerstone of energy cooperation in the eastern Mediterranean.
El-Molla said that the fact that Cairo is the permanent host for the newly found forum, confirms its centrality as a regional centre for trading of gas and oil.
Ellinas noted that Egypt is becoming a major player in its own right. "Not only it has become self-sufficient in gas, but it is fast promoting renewables and by removing subsidies it is containing its growing domestic energy demand. This has freed gas for exports from its existing liquefaction facilities. With increasing gas discoveries and production, it expects these to reach full utilisation be next year," he said, adding that some gas from its neighbours, Israel and Cyprus, may reach Egypt if prices make it commercially viable.
Russia, Turkey mull joint exploration in Mediterranean
Meanwhile, Russian oil companies could roll out exploration in offshore Mediterranean in cooperation with Turkey, Anadolu news agency quoted Russian Energy Minister Alexander Novak as saying in an interview on 26 July. Russian state oil giant Rosneft is working at Zohr gas field in Egypt.
Asked if Moscow is trying to involve Ankara in order to get a foothold in the region, Ellinas said at present such cooperation is for exploration in Turkish waters. "It is difficult to see how this can provide Russia with a foothold in the region, if indeed it wants to. Russian companies had chances to complete for blocks in Cyprus EEZ but did not use them," he said, adding "And (Russian gas producer) Novatek is in Lebanon already."
Source: neweurope.eu


text:
THQ Nordic Now Owns the Outcast IP
THQ Nordic has been on something of a spending spree over the last 12 months, and the latest acquisition has just been sent through via press release.
THQ Nordic now owns the Outcast IP after having agreed a deal with the original creators of the game. The game originally released way back in 1999 before being remade and re-released as Outcast – Second Contact modern systems where it received fairly positive reviews.
Now that THQ Nordic owns the game and its rights, it's possible we may get more sequels in the not-so-distant future…
We sometimes link to online retail stores. If you buy something from our links, we may make a small commission which goes towards keeping the lights on and coffee in the pot.
Chris has been writing about gaming news for far too long, and now he's doing it even more. A true PlayStation know-it-all, Chris has owned just about every Sony console that ever existed. Trophies are like crack to this fella. (Bronze trophies, that is – he only has one Platinum.)

subset: commoncrawl


text:
Disk Read Error Press Ctrl Alt Del To Recover
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text:
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Mikael is a Scandinavian form of the biblical name Michael.
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Vossoughi, Shirin; Jackson, Ava; Chen, Suzanne; Roldan, Wendy; Escudé, Meg – Journal of the Learning Sciences, 2020
Studies of embodied cognition offer powerful accounts of the semiotic resources people use as they think together within different domains. Yet this research does not typically foreground the history of relationships within focal interactions--a history we have found to be consequential to the ways embodied actions unfold. Through ethnographic and…
Descriptors: Cognitive Processes, Human Body, Ethics, Relationship
Ramey, Kay E.; Uttal, David H. – Journal of the Learning Sciences, 2017
Spatial thinking is important for success in engineering. However, little is known about "how" students learn and apply spatial skills, particularly in kindergarten to Grade 12 engineering learning. The present study investigated the role of spatial thinking in engineering learning at a middle school summer camp. Participants were 26…
Descriptors: Summer Programs, Middle School Students, Spatial Ability, Engineering Education
Ottmar, Erin; Landy, David – Journal of the Learning Sciences, 2017
Learning algebra is difficult for many students in part because of an emphasis on the memorization of abstract rules. Algebraic reasoners across expertise levels often rely on perceptual-motor strategies to make these rules meaningful and memorable. However, in many cases, rules are provided as patterns to be memorized verbally, with little overt…
Descriptors: Mathematics Instruction, Algebra, Outcomes of Education, Learning Processes
Headrick Taylor, Katie – Journal of the Learning Sciences, 2017
The "everyware" paradigm opens up new possibilities for learning on-the-move with technologies through urban spaces while also raising questions about emerging literacies required of users to understand and use the digital traces these technologies generate. This article develops locative literacies as a way of understanding place-based,…
Descriptors: Urban Areas, Geographic Information Systems, Geographic Location, Navigation
Kapur, Manu – Journal of the Learning Sciences, 2014
A total of 136 eighth-grade math students from 2 Singapore schools learned from either productive failure (PF) or vicarious failure (VF). PF students "generated" solutions to a complex problem targeting the concept of variance that they had not learned yet before receiving instruction on the targeted concept. VF students…
Descriptors: Foreign Countries, Grade 8, Mathematics Instruction, Failure
Jeppsson, Fredrik; Haglund, Jesper; Amin, Tamer G.; Stromdahl, Helge – Journal of the Learning Sciences, 2013
A growing body of research has examined the experiential grounding of scientific thought and the role of experiential intuitive knowledge in science learning. Meanwhile, research in cognitive linguistics has identified many "conceptual metaphors" (CMs), metaphorical mappings between abstract concepts and experiential source domains,…
Descriptors: Abstract Reasoning, Chemistry, Figurative Language, Cognitive Processes
Abrahamson, Dor – Journal of the Learning Sciences, 2012
Some intensive quantities, such as slope, velocity, or likelihood, are perceptually privileged in the sense that they are experienced as holistic, irreducible sensations. However, the formal expression of these quantities uses "a/b" analytic metrics; for example, the slope of a line is the quotient of its rise and run. Thus, whereas students'…
Descriptors: Teaching Methods, Mathematics Instruction, Middle School Students, Thinking Skills
Reed, Stephen K. – Journal of the Learning Sciences, 2012
Many types of learning require the mapping of information across situations. The proposed organizational framework extends the cognitive study of mappings across problems to include mappings across representations, solutions, and sociocultural contexts. I apply one-to-one, one-to-many, and partial mappings to analyze representative cases that…
Descriptors: Concept Formation, Cognitive Processes, Logical Thinking, Information Transfer
Lobato, Joanne; Rhodehamel, Bohdan; Hohensee, Charles – Journal of the Learning Sciences, 2012
As transfer researchers have begun to investigate a broader range of phenomena, they have correspondingly put forward new processes to provide explanatory accounts for the occurrence of transfer. This move coincides with a call to acknowledge the contribution of social interactions, language, cultural artifacts, and normed practices to the…
Descriptors: Transfer of Training, Mathematics Instruction, Cognitive Processes, Interpersonal Communication
On the Science of Embodied Cognition in the 2010s: Research Questions, Appropriate Reductionism, and Testable Explanations
Nunez, Rafael – Journal of the Learning Sciences, 2012
"The Journal of the Learning Sciences" has devoted this special issue to the study of embodied cognition (as it applies to mathematics), a topic that for several decades has gained attention in the cognitive sciences and in mathematics education, in particular. In this commentary, the author aims to address crucial questions in embodied…
Descriptors: Mathematics Education, Cognitive Processes, Human Body, Holistic Approach
Stevens, Reed – Journal of the Learning Sciences, 2012
For a long time, the body has had a bad rap among some big names in the Western philosophical tradition. Plato and Descartes come quickly to mind as particularly bad rappers. But it appears that the body is making a comeback in the cognitive and learning sciences. Long banished from the main stage by an idealized, inside-the-head…
Descriptors: Phenomenology, Human Body, Cognitive Processes, Holistic Approach
When the Classroom Floor Becomes the Complex Plane: Addition and Multiplication as Ways of Bodily Navigation
Nemirovsky, Ricardo; Rasmussen, Chris; Sweeney, George; Wawro, Megan – Journal of the Learning Sciences, 2012
In this article we contribute a perspective on mathematical embodied cognition consistent with a phenomenological understanding of perception and body motion. It is based on the analysis of 4 selected episodes in 1 session of an undergraduate mathematics class. The theme of this particular class session was the geometric interpretation of the…
Descriptors: Cognitive Processes, Mathematics Education, Perception, Mobility
Alibali, Martha W.; Nathan, Mitchell J. – Journal of the Learning Sciences, 2012
Gestures are often taken as evidence that the body is involved in thinking and speaking about the ideas expressed in those gestures. In this article, we present evidence drawn from teachers' and learners' gestures to make the case that mathematical knowledge is embodied. We argue that mathematical cognition is embodied in 2 key senses: It is based…
Descriptors: Mathematics Education, Mathematical Concepts, Physical Environment, Mathematics Instruction
Design and Reflection Help Students Develop Scientific Abilities: Learning in Introductory Physics Laboratories
Etkina, Eugenia; Karelina, Anna; Ruibal-Villasenor, Maria; Rosengrant, David; Jordan, Rebecca; Hmelo-Silver, Cindy E. – Journal of the Learning Sciences, 2010
Design activities, when embedded in an inquiry cycle and appropriately scaffolded and supplemented with reflection, can promote the development of the habits of mind (scientific abilities) that are an important part of scientific practice. Through the Investigative Science Learning Environment ("ISLE"), students construct physics knowledge by…
Descriptors: Science Process Skills, Skill Development, Cognitive Processes, Apprenticeships
Wagner, Joseph F. – Journal of the Learning Sciences, 2010
Many approaches to the transfer problem argue that transfer depends on the recognition of the same or similar abstract "structure" in 2 different situations. However, mainstream cognitive perspectives and contrasting Piagetian constructivist accounts differ in their conceptualizations of structure. These differences, not clearly articulated in the…
Descriptors: Transfer of Training, Constructivism (Learning), Undergraduate Students, Mathematical Concepts
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Exams Manager
Activate Learning is the largest provider of further education and training in Oxfordshire and one of the leading colleges in the learning & skills sector. We are a forward thinking, commercially focussed inspirational place to work and learn. With high levels of learner achievement and significant investment in resources.
The Examinations Team are responsible for all matters relating to the students' outcomes – obtaining information from Faculties, registration of students with awarding bodies, entering students for exams, running of the exams, processing of results and providing examination data as required. An exciting opportunity has arisen for an Exams Manager (Oxfordshire) to join our busy team. The core role involves managing and leading the examinations officers and supporting the smooth running of all examination processes within our Oxfordshire College sites (City of Oxford College, Technology Campus in Blackbird Leys and Banbury & Bicester College.
You will have excellent organisational and teamwork skills, and a high level of customer service skills to enable you to communicate effectively with all our customers. Previous experience and confidence in using a large data base is essential. Strong IT skills and the ability to use all Microsoft programmes including Outlook, Excel and Word are essential. Management experience within an educational environment is desirable, as the ability to manage and lead a team across sites will be a core role. The ability to use your own initiative, and be friendly and confident in dealing with day to day systems, procedures and students is essential.
Travel across our Oxfordshire sites is a requirement for this role.
If you like a challenge, then this is the post for you. We look forward to meeting you.
Interviews will take place at the City of Oxford College, on Thursday 31st October 2019
To apply, please complete an application form or attach a CV and send to the email address below, with 'College Jobs' as the subject.
- Recruiter
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text:
Villa Palladio is an elegant villa near Siena, sleeps 14
Back to the Villas for
10 - 16
Villa Palladio is an elegant 18th century mansion near Siena in the Chianti wine region, The villa was recently renovated, enhancing the charm of the villa without altering its original structure, Today, Villa Palladio is the perfect place to enjoy a pleasant stay under the Tuscan sun, ideally situated to visit Tuscany's most important historical and cultural cities. The villa was originally used by noble families as a relaxing place to escape to in the countryside during the hottest months of summer.
Its position in the centre of Tuscany, just a few kms from the main Firenze-Siena road and near the historic intersection between the Via Cassia and Via Francigena, makes this villa the perfect location to explore all of Tuscany - from wine roads to visiting Tuscany's many cultural and artistic sites, not forgetting the opportunity to sample good Tuscan cuisine in the many traditional villages of the area.
|V. High
|1 April -
11 May
13 Oct - 20 Dec
|11 May -
31 May
1 Oct - 13 Oct
20 Dec - 10 Jan/20
|1 June -
30 June
1 Sept - 30 Sept
|1 July - 31 Aug
Rental: Weekly, Sat to Sat
Non Sat arrival between 1 April - 11 May, 13 Oct - 20 Dec, 20 Dec - 10 Jan
3 nt stay between 1 April - 11 May, 13 Oct - 20 Dec
Villa closed between 10 Jan - 31 March
Prices include: Water, kitchen gas, electricity up to 400kw, maid service mid week (Wed), linen & towels (changed weekly), WI FI internet, toiletries (shampoo, soap, hairdryer, toilet paper)
Not included: Extra maid service - €15 per hr, extra bed - €125 per week, extra linen and towel change: bathmat €2, guest towel €3, hand towel €3, bath towel €4, pool towel (not provided and available on request) €5, bathrobe (not provided and available on request) €8, single sheet set €7, Queen\King sheet set €10, pillowcase €2, phone/fax, heating €4,50 per unit, electricity over 400kw – €0,40 per unit, first grocery shopping service €30 plus cost of food, breakfast €15 per person, €10 under 12 yrs, final cleaning - €180
Tourist Tax: €1.50 per adult per night, payable in cash on arrival
Security Deposit: €500 in cash on arrival, returned on departure in full or as agreed
The ground floor is typical of an 18th century villa inspired by the Palladian style. On the right side of the large entrance is a wide, bright living room, which is the largest and oldest section of the house (120 sq.m.). Characterized by its vaults, the living room is enhanced by a unique fireplace in green marble and is furnished with antique furniture. Off the livingroom is a double bedroom with ensuite bathroom. From the livingroom, a glass door leads outside to the sunny terrace where there are several lemon plants.
On the left side of the entrance, there is an elegant well-lit dining room with three windows overlooking the Villa gardens. From the dining room you can access the kitchen, which has been completely renovated and has a wide marble counter-top which is ideal for food preparation, baking etc. The kitchen is fully equipped and has an independent service entrance behind the villa.
All bedrooms on the first floor sleeping area are fully equipped with air conditioning.The staircase from the ground floor opens onto a grand living room with access to a balcony where you can enjoy a wonderful view of Monteriggioni. This living room leads to the first of three double bedrooms with en-suite bathrooms. A large, bright hallway leads to the other three bedrooms, two twin bedrooms with shared bathroom and one double bedroom with en-suite bathroom.
The villa is surrounded by a fully fenced 1,000 sqm garden. On the large area in front of the villa is a beautiful gazebo suitable for outdoor dining. To the side of the villa, guests can enjoy relaxing on the terrace where there are lemon trees and other shaded areas
In a secluded area to the back of the villa, nestled among cypress and olive trees, is the swimming pool surrounded by a large lawn. The pool is 10 x 5m and has deckchairs and sun umbrellas.
Private parking
Large garden
Private pool 10x5 mts May - Oct
Outdoor dining
Well equipped kitchen
Washing machine
WI FI internet
Multifunction printer
Air con in bedrooms
Check in 3pm - 7pm
Check out by 10am
Local village - 2km
Monteriggioni - 5km
Poggibonsi - 8km
Siena - 15km
San Gimignano - 20km
Florence - 50km
Pisa airport - 90km


text:
Shaunti Feldhahn
Today, she applies her analytical skills to investigating eye-opening, life-changing truths about relationships, both at home and in the workplace.
Shaunti's "Find Series" of devotionals and Bible studies includes Find Rest, Find Balance, and Find Peace – and guides women to discover the path that will take them from frenzied to peace when they lean into God's truths and begin to live the life designed just for them.
Shaunti's findings have been featured in media as diverse as The Today Show and Focus on the Family, The New York Times and iMom. Shaunti lives in Atlanta with her husband, Jeff, their two teenage children, and two cats who think they are dogs.
Find Rest
Women often find themselves exhausted by their demanding schedules and endless commitments. Shaunti brings us down the path to abundant peace through her brand-new 60-day devotional, Find Rest..
Find Peace
In this daily devotional, we'll discover the biblical direction to become a woman of serenity and delight in all seasons and have impact for generations to come.
Find Balance
From stay-at-home moms to globe-trotting executives, every modern woman has asked, How do I juggle it all? Now, Find Balance helps you discover God's road map to peace, place, rest, and delight in this crazy, contemporary life.
Find Rest
With its scripture reminders and quotes, this journal is the perfect way to spend quiet time with God, remembering His biblical promises, and reflecting on your journey to find rest in the middle of a busy life.
Shaunti Feldhahn
Complete with scripture and quotes, this journal will help and encourage women at any stage as they reflect on their motherhood journey. Remembering God's promises and truths, even amidst the uncertainty and fear of parenting, will provide the joy, purpose, and peace only Jesus has to offer.
Find Balance
This special edition format of Find Balance includes two beautiful art prints and the Find Balance study, all packaged in a durable, patterned sleeve.


text:
Experimental investigation on high strength experimental investigation on high strength concrete as partial replacement of natural sand by manufactured sand and quarry dust e suresh 1 i bargathulla 2 1pg student new prince shri bhavani college of engineering and technology chennai tamil n adu india
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Experimental investigation on high strength experimental investigation on high strength concrete as partial replacement of natural sand by manufactured sand and quarry dust e suresh 1 i bargathulla 2 1pg student new prince shri bhavani college of engineering and technology chennai tamil n adu indiacome look
The paper investigated the application of gravel as partial economic replacement of granite in concrete production concrete was produced using granitegravel combination in varying percentages of 9010 8020 7030 6040 5050 4060 3070 2080 and 1090 concrete made from 100 granite and 100 gravel served controls while other constituents of concrete were kept constantcome look
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We previously demonstrated that mannoprotein (MP) from Cryptococcus neoformans (CnMP) stimulates interleukin-12 production by human monocytes, thus fostering a T-helper type 1 (Th1) protective anticryptococcal response. In this paper we show that CnMP was also able to induce a Candida albicans-directed protective Th1 response. This was demonstrated for mice immunized with CnMP by induction of a delayed-type hypersensitivity (DTH) reaction to C. albicans MP (CaMP) as well as induction of gamma interferon production by CD4+ and CD8+ splenic T cells stimulated in vitro with CaMP. CnMP-immunized mice were also partially protected from lethal systemic challenge with C. albicans, as shown by prolonged median survival times and decreased fungal burden in the kidney. Much evidence supports the validity of these cross-reactive and functional Th1 responses: (i) a non-cross-reactive C. albicans antigen, such as enolase, did not produce a DTH response to CaMP; (ii) passive adoptive transfer of T cells primed with CnMP induced a DTH reaction; (iii) C. neoformans extract elicited a DTH response to CaMP; and (iv) a monoclonal antibody (7H6) directed against a major and immunodominant T-cell-stimulatory 65-kDa MP (MP65) of C. albicans also recognized discrete 100-kDa constituents of C. neoformans extracts, as well as secretory constituents of the fungus. These results suggest the presence of common Th1 antigenic determinants in the mannoproteic material of C. neoformans and C. albicans epitopes, which should be considered in devising common strategies for immunoprophylactic or immunotherapeutic control of the fungi.
Cryptococcus neoformans and Candida albicans are common causes of opportunistic fungal disease in immunocompromised or otherwise modified hosts (1). Both innate and adaptive immune responses have been shown elsewhere to contribute to host resistance to candidiasis (24), while a complex interplay between humoral and cellular immunity is critical for the control of cryptococcal infection (31).
Mannoprotein (MP) is a natural glycoconjugate usually containing between 80 and 90% mannose expressed mainly on the fungal surface and released into the external medium during growth. MPs are abundant in the cell wall of C. albicans and constitute about one-third of the dry mass (3). One of them, an immunodominant 65-kDa-MP antigen (MP65), has been extensively characterized both biochemically and immunologically (7, 8). Other MPs have been described as enzymes or adhesins involved in the pathogenicity of this fungus (10). The capsular material of C. neoformans is mainly composed of glucuronoxylomannan (galactose and xylose are minor constituents), whereas MPs are minor components of the capsule and the cell wall (6, 30).
For both organisms, MPs have long been recognized as important antigens involved in the induction of the T-cell-mediated immune response, i.e., a critical response for antifungal protection in humans. In particular, Candida and Cryptococcus MPs were shown elsewhere to induce lymphoproliferation (19, 23) and cytokine production (21, 22, 27). Previous data showed that interleukin-12 (IL-12) plays a pivotal role in induction of the T-helper type 1 (Th1) response against C. neoformans infection, this response being essential for protection (6). More recently it was demonstrated that an MP preparation of C. neoformans induces early (3 to 6 h) production of IL-12 by human monocytes in vitro, resulting in early (12 to 24 h) secretion of gamma interferon (IFN-γ) by T cells (22). Furthermore, in vivo treatment with MP induces IL-12 secretion by splenic macrophages and IL-12 p40 mRNA expression in the brain. MP-induced IL-12 and IFN-γ secretion coincides with enhanced antifungal activity, resolution of the inflammatory process, and clearance of fungal load from the brain (21).
Recent data on human monocytes stimulated in vitro by yeast or mycelial forms of C. albicans also suggest that surface-expressed MP may differentially induce IL-12, up- or down-regulating a Th1 protective cytokine pattern (5; unpublished data). When specifically examined, stimulatory properties of MPs in cell-mediated immunity (CMI) were found to be expressed by the protein rather than by the saccharide moiety of these molecules (7, 8, 18).
Because of these interesting properties, further biochemical and immunological characterization of MPs from C. albicans and C. neoformans could be extremely useful in devising immunotherapeutic or vaccination strategies against both fungi. We were particularly attracted by the possibility that protein epitopes common to C. neoformans and C. albicans MPs are involved in the induction of protective CMI. For this paper we examined this hypothesis by immunizing mice with C. neoformans MP (CnMP) and assessing C. albicans MP (CaMP)-specific CMI responses and protection from C. albicans challenge in mice.
Mice.Female CD1 mice purchased from Harlan Italy Laboratories (Udine, Italy) were used at 4 to 6 weeks of age.
Microorganisms.A virulent germ-tube-forming strain of C. albicans (CA-6) isolated from a clinical specimen was used in this study. The origin, characteristics, and growth conditions of CA-6 have been described previously (2). An agerminative strain of C. albicans (PCA-2) was kindly supplied by D. Kerridge (Department of Biochemistry, University of Cambridge, Cambridge, United Kingdom). This strain grows as a pure yeast form in vitro at 28 or at 37°C in conventional mycological media.
C. neoformans strain 6995, a thinly encapsulated strain of serotype A from the Centraalbureau voor Schimmelcultures, Delft, The Netherlands (CBS 6995 = NIH 37), was also used in this study. The cultures were maintained by serial passage on Sabouraud agar (BioMérieux, Lyon, France). In selected experiments an acapsular mutant of C. neoformans CAP67 obtained from the American Type Culture Collection (Manassas, Va.) was used. Log-phase yeasts were harvested by suspending a single colony in saline; cells were then counted on a hemocytometer and adjusted to the desired concentration. Yeasts were killed by being heated at 60°C for 30 min.
Preparation of MP extracts of C. neoformans.An acapsular mutant of C. neoformans (NIH B-4131) was cultured in a defined medium containing 2% glucose for 5 days at 35°C as previously described (9). The culture supernatant containing MP was concentrated by ultrafiltration, and purification was performed by a combination of affinity chromatography (concanavalin A [ConA]) and anion-exchange chromatography (DEAE) (Whatman; Chemical Separation Ltd., Maidstone, England) (29). Chemically, CnMP extract used in this study consists of 13.3% protein, 72% carbohydrate, and 3.1% hexosamine (4). CnMP did not contain any contaminating ConA as analyzed by sodium dodecyl phosphate-polyacrylamide gel electrophoresis, amino acid analysis, and molecular mass determination (ConA has a molecular mass of 102 kDa). Intraperitoneal treatment with ConA (5 mg/kg of body weight, in 200 μl of saline) 6 and 24 h before C. albicans challenge did not affect survival of mice challenged with lethal doses of C. albicans or produce a delayed-type hypersensitivity (DTH) response to CaMP or CnMP challenge. This indicates that MPs had no contaminating biologically active ConA.
Preparation of MP extracts and enolase of C. albicans.A crude MP extract was obtained from washed yeast-form C. albicans BP serotype A. MP-F2, a major MP fraction of the MP extract, was subsequently separated by ion-exchange chromatography on a DEAE Sephadex A-50 column. Only one defined batch of MP-F2 (identified as CaMP) was used for all experiments. The preparation was passed through a Sepharose-polymyxin B column (Sigma, Milan, Italy) to ensure the absence of contamination by bacterial endotoxin. A full description of the basic chemical and molecular composition of the MP-F2 fraction has been previously published (20, 26). Recombinant enolase generated as previously described (25) was used in this study as control MP-nonrelated C. albicans antigen.
Maintenance of endotoxin-free conditions.Preparations of the various cryptococcal and candidacidal components tested negative for endotoxin contamination by a Limulus assay (Coatest endotoxin; Kabi Diagnostica, Mölndal, Sweden) with a sensitivity of 25 pg of Escherichia coli lipopolysaccharide. Nevertheless, all in vitro experiments were carried out at least once in the presence of 10 μg of polymyxin B (Sigma) per ml to neutralize any undetected contamination with bacterial lipopolysaccharide.
MAbs.The monoclonal antibody (MAb) 7H6, which recognizes a peptide epitope of MP65, has been described previously (7, 8). It was used in immunoblots as described below. Monoclonal rat anti-mouse CD3 was obtained from BD Pharmingen (Heidelberg, Germany).
Sodium dodecyl sulfate-polyacrylamide gel electrophoresis and Western blotting.Sodium dodecyl sulfate-polyacrylamide gel electrophoresis was performed by the Laemmli method (11) with a minigel system (mini-Protean II; Bio-Rad Laboratories, Hercules, Calif.). Samples were separated in 12% polyacrylamide gels with a 4% acrylamide stacking gel at a constant voltage of 180 V for 60 min. After electrophoresis, MPs were stained with the Bio-Rad silver staining kit or transferred electrophoretically onto polyvinylidene difluoride membranes in a Trans Blot cell (Bio-Rad). After periodate treatment, membranes were reacted with MAb 7H6. Purified MAb was used at a 1:50 dilution and detected with peroxidase-conjugated anti-mouse immunoglobulins (Boehringer Mannheim GmbH, Mannheim, Germany) at a 1:250 dilution.
Infection with C. albicans. C. albicans yeasts from a 24-h culture on Sabouraud agar were washed with sterile endotoxin-free physiological saline (Fresenius Kabi Italia SpA, Verona, Italy); counts were determined on a hemocytometer, and the suspension was adjusted to the desired concentration for intravenous injection. The number of viable yeast cells injected was confirmed by culturing dilutions of the inoculum on Sabouraud agar. Infection was accomplished by injecting 0.5 ml of prewarmed yeast cell suspension (5 × 105 yeast cells/ml) through a 27-gauge needle into the tail vein. The outcome of infection was evaluated in terms of median survival time, number of dead animals over total number of animals, and fungal burden in the kidney expressed as CFU.
In vivo treatment.Groups of 7 to 10 mice were injected intraperitoneally with 10 μg of CnMP in saline 24 and 6 h before infection. Mice injected with saline alone served as controls.
Clearance of Candida from kidneys.At designated times after infection mice were sacrificed and kidneys were removed and homogenized. A serial 10-fold dilution of each sample was plated in duplicate on Sabouraud agar plates and incubated for 24 h, and CFU were counted.
Preparation and stimulation of spleen cells.After MP treatment, spleens were aseptically removed and placed in 5 ml of RPMI 1640. Splenocytes were cultured to analyze cytokine production. A total of 20 × 106 cells/ml were cultured alone or in the presence of CnMP or CaMP (2.5 μg/ml), heat-inactivated C. neoformans (20 × 106 cells/ml), or heat-inactivated C. albicans (20 × 106 cells/ml) in RPMI-10% fetal calf serum. As a positive control, cells were stimulated with 50 ng of phorbol 12-myristate 13-acetate (PMA; Sigma)/ml and 2 mM ionomycin (Sigma). After 18 h of incubation, intracellular IFN-γ was measured. In parallel experiments, after 7 days the supernatants were assayed for IFN-γ content.
Determination of IFN-γ production.Cytokine levels in culture supernatant fluids were measured with an enzyme-linked immunosorbent assay kit for mouse IFN-γ (Endogen Inc., Woburn, Mass.).
Intracellular IFN-γ assay.The intracellular IFN-γ in CD4 or CD8 T cells was determined with an intracellular staining kit (IC Screen) provided by Biosource International (Camarillo, Calif.). Stimulations were carried out in the presence of the transport inhibitor monensin, which prevents the release of IFN-γ into the extracellular milieu. The stimulated cells were stained with fluorescein isothiocyanate-tagged anti-CD4 or anti-CD8 MAb and fixed with paraformaldehyde. Cells were stained with an R-phycoerythrin-labeled anti-IFN-γ antibody in the presence of a permeabilizing agent. The stained cells were then analyzed by flow cytometry for the coexistence of two different fluorescent tags in the same cell.
Induction and elicitation of the DTH response.Mice were intraperitoneally injected twice in 24 h with 200 μl of saline or 10 μg of CnMP in saline. After 8 days the left footpads were injected with 30 μl (10 μg) of CnMP, CaMP, or extracts from the acapsular strain (7698) and the right footpad was injected with 30 μl of saline. The DTH reaction was recorded 24 h later by weighing the footpads as a measure of swelling, and the results were expressed as the increase in weight of the left hind footpad over that of the saline-injected right hind counterpart. Data were expressed as means ± standard errors of the means (SEM) of five mice per group per experiment.
Adoptive transfer of T cells.Mice were injected with 10 μg of CnMP twice in 24 h. T cells were purified from spleens as described previously (15) 7 days after treatment and restimulated in vitro with CnMP for 24 h. These cells (more than 95% CD3 positive as revealed by flow cytometric analysis) were injected at the dose of 106 into naïve mice that had been challenged in the footpad with CaMP or saline. The DTH reaction was recorded 24 h later.
Statistical analysis.Statistical significance between groups was analyzed by the Mann-Whitney U test. Survival of the different groups of mice in the protection experiments was compared by the log-rank survival test. For in vitro experiments statistical analysis was performed by the analysis of variance test. A P value of <0.05 was considered significant.
In consideration of the previously published evidence that both CnMP and CaMP could induce and detect specific CMI responses (16, 17), we explored the possibility of a cross-reactive antigenic response to the two fungi by immunizing mice with CnMP and then assessing DTH reaction and IFN-γ production by splenic lymphocytes stimulated in vitro with CnMP or CaMP.
As expected, mice immunized with CnMP generated a strong DTH reaction when the footpad was challenged 7 days posttreatment with the corresponding antigen, compared to that of untreated control mice. Interestingly, a marked footpad swelling was also detected in mice injected with material from C. neoformans, such as 7698 extracts and CnMP and DTH tested with CaMP (Fig. 1). The difference between the two reactions (against CnMP and CaMP) was not statistically significant. The reaction specificity was confirmed by the fact that a non-cross-reactive antigen, enolase, did not induce a DTH response when used instead of CaMP.
The cross-reactive immune response between CaMP and CnMP was also demonstrated by the fact that reverse treatment with CaMP induced a DTH response against CnMP similar to that obtained with the corresponding antigen (CnMP) (Fig. 1).
To verify whether T cells primed with CnMP were indeed responsible for DTH induction, splenic T cells (106) from naïve or CnMP-treated mice (6 days posttreatment) were adoptively transferred intravenously into naïve mice 1 day before CaMP challenge. The results showed that T cells from CnMP-treated mice produced a significant increase of 5.6 ± 0.7 mg in footpad weight (P < 0.05, CnMP treated versus saline treated). Conversely, a nonsignificant difference was observed in mice receiving T cells from naïve mice (increase of footpad weight was 1.9 ± 0.5 mg).
Since IFN-γ is critically involved in the generation and expression of the DTH reaction, we examined whether, and to what extent, this cytokine was produced in the supernatants of splenocyte cultures from mice immunized with CnMP and stimulated in vitro with CnMP or CaMP or other control stimuli. Figure 2 shows that, as early as 1 day after in vitro stimulation, splenocytes from CnMP-immunized animals, but not from nonimmune mice, produced elevated quantities of the cytokine (from about 150 to more than 450 pg/ml) in response to in vitro treatments not only with the specific immunogens but also with CaMP and whole inactivated C. albicans cells (PCA-2). Interestingly, in vitro stimulation with whole cells of the fungus was the same, if not superior, in terms of cytokine production as stimulation of whole C. neoformans (6995) (Fig. 2). In addition, appreciable levels of IFN-γ were observed upon stimulation with extracts from C. neoformans cells (7698). Conversely, a non-cross-reactive unrelated antigen, enolase, did not induce IFN-γ (Fig. 2). A substantial drop in IFN-γ production (no more than 25 pg/ml) was observed when the cytokine was sought in splenocyte supernatant cultures from mice treated 7 days before with CnMP. In this case, no significant difference was found between the low, spontaneous cytokine production of unstimulated cells and that of the cells stimulated by C. neoformans or C. albicans materials (data not shown)
Knowing that IFN-γ may be secreted by CD4 or CD8 T cells and that CD8 could also be involved in antifungal protection (1), we determined the percentage of IFN-γ-producing CD4 and CD8 cells over the total number of splenic lymphocytes. As shown in Fig. 3, a substantial proportion (approximately 15 to 20%) of both CD4 and CD8 splenic lymphocytes of mice immunized with CnMP produced IFN-γ in the presence of both cryptococcal and candidal materials. In particular, this production was not inferior to that of the positive control with PMA plus ionomycin-treated cells. On average, about 5% of the CD8 T cells from immune animals were IFN-γ positive and the percentage increased about threefold upon in vitro stimulation (Fig. 3). No statistically significant difference was found in the percentage of CD4 or CD8 cells positive for intracellular IFN-γ among the various cultures stimulated with cryptococcal or candidal materials. Even though the percentages of cytokine-producing CD4 and CD8 cells were similar, it is likely that the major contribution to IFN-γ production may be ascribed to CD4+ cells because they represent more than 65% of total T cells among splenocytes of CnMP-treated mice.
Given that CnMP is able to induce a specific T-cell response to CaMP, and the major antigenic component of CaMP capable of inducing a T-cell response is a 65-kDa MP (MP65) (7), we hypothesized that CnMP could share common epitopes with MP65. Thus, we verified the ability of a MAb specific for a protein epitope of the N-terminal moiety of MP65 (MAb 7H6) (A. Cassone, unpublished observations) to recognize CnMP and/or other cryptococcal components present in cellular extracts or secreted by the fungus. Experiments with CnMP showed no reaction with MAb 7H6. However, other extracts of C. neoformans and CaMP were indeed recognized by the antibody (Fig. 4). C. neoformans extract demonstrated a well-defined, discrete reactivity with a molecular constituent of about 100 kDa, both for the acapsular and for the encapsulated fungus. Reactivity with the secretory materials of the encapsulated strain (6995) was less defined and more polydisperse than in the extract (Fig. 4).
Finally, in view of the report that treatment with selected MP fractions of C. albicans induced partial protection against systemic infection by this fungus (owing to the activation of a Th1-type response) (16), we examined mice immunized with CnMP to determine if they were also able to resist systemic challenge by C. albicans. To this end, the animals were injected intraperitoneally with CnMP (10 μg/200 μl) 24 and 6 h before intravenous challenge with C. albicans (2.5 × 105 cells/0.5 ml), and animal survival and fungal CFU were determined. As shown in Fig. 5A, survival of mice immunized with CnMP was significantly higher than that of nonimmune animals, corresponding to the statistically significant reduction of the Candida burden in the kidney (Fig. 5B). Similar results in terms of survival were obtained in our experimental system by treatment with the corresponding antigen CaMP (Fig. 5A).
The protective effect of CnMP was also evaluated against the homologous strain of C. neoformans (CAP67). Given that acapsular C. neoformans CAP67 is unable to produce lethal infection in mice, we evaluated CFU from brains 7 days after CAP67 (5 × 106) intravenous challenge. The results showed that CnMP treatment produced a reduction of C. neoformans load in the brain as evaluated by CFU recovery (data not shown).
The results reported here demonstrate that MPs from C. neoformans and C. albicans share common antigenic determinants capable of eliciting potentially protective Th1-type responses in immunized mice. This assertion derives from three lines of evidence. First, animals primed with CnMP generated a DTH response against CaMP to a degree that did not differ substantially from the response to the priming antigen CnMP. This response was immune mediated as shown by T-cell adoptive transfer. Second, splenocytes of mice immunized with CnMP produced IFN-γ when cultured in vitro with CaMP. Third, IFN-γ-producing CD4 and CD8 T cells were markedly increased in animals immunized with CnMP. CD4 and CD8 cell expansion upon in vitro stimulation with CaMP was comparable to that observed with the priming antigen and also similar to that detected with whole C. albicans and C. neoformans as stimulants. Finally, the cross-reactive T-cell response was functional in terms of protection against C. albicans and C. neoformans.
As a separate yet connected piece of evidence for the sharing of antigenic MP determinants between the two fungi, we show here that MAb 7H6 recognizes a peptide epitope close to the N-terminal moiety of a major immunodominant T-cell immunogen of C. albicans MP65 (7; A. Cassone, unpublished data) and reacts with defined molecular constituents of C. neoformans extracts, suggesting that a homologous molecular constituent may be present in CnMPs. This is consistent with the ability of C. neoformans extracts to elicit a specific DTH response against C. albicans. The lack of reaction of MAb 7H6 with CnMP may not necessarily predict the absence of a MAb-reactive epitope also in CnMP, as it could have become unreactive during the process of CnMP preparation, which includes heating at 100°C for 5 min. If so, this epitope may be present in a molecule that is smaller than C. albicans MP65, since the calculated molecular mass of CnMP is about 8.2 kDa (4). Our data, in particular the strong reaction of MAb 7H6 with a 75-kDa constituent of C. neoformans extract, may also imply that CnMP is a degradation product of this or another polydisperse MAb-reactive extract or secretory material of C. neoformans. Studies of the molecular cloning of the MP65-encoding gene of C. albicans are in progress (12).
MP molecules have long been argued to be important constituents for the pathogen-host relationship in both C. neoformans and C. albicans. MP65 of C. albicans has been shown elsewhere to possess strong immunogenic properties, including DTH elicitation (16) and induction of T-cell responses (23). It has been recognized to contain dominant T-cell epitopes in humans (12) and is used to expand specific T cells, in particular CD4+ T-cell clones producing IFN-γ (18). Partial protection against systemic challenge by C. albicans in mice has also been attributed to an anti-MP65 response (16).
MPs of C. neoformans have been identified elsewhere as the major antigens inducing CMI responses to C. neoformans (17), including DTH elicitation, T-cell activation, and cytokine production by peripheral blood mononuclear cells (4, 19, 22). More recently the molecular characterization of a C. neoformans MP that stimulates a T-cell response has been provided (13, 14). We have shown that our MP preparation from C. neoformans is capable of inducing IL-12 production from human monocytes and protection against a lethal fungus challenge (21, 22).
Thus, MPs from C. albicans and C. neoformans appear to play an important role in the elicitation of protective antifungal responses. The data provided in this paper not only strengthen this concept but also suggest the involvement of a common determinant in these molecules. The specificity of the T-cell response has been underlined by the inability of MP to affect the blastogenic response to mitogens, such as ConA. More evidence on the specific immune response elicited by MP from C. neoformans has been provided by the induction of IFN-γ secretion from CD4 and CD8 T cells. It is noteworthy that there is a profound difference between IFN-γ recovered after 1 day and that recovered after 7 days. Peak IFN-γ production occurring as early as 1 day after stimulation is consistent with the fact that MP induces an early and massive production of IL-12 after 6 h (22). Hence, the abundant production after 1 day of culture likely reflects this event, since IL-12 and IFN-γ production are closely related (28). This is consistent with the fact that MP from C. albicans stimulates CD4 T cells (18) and that CD4 and CD8 lymphocytes are important in controlling C. albicans infection (1). The strategy of the immune system in mounting a specific response against C. albicans indicates that MP from C. neoformans contains common epitopes potentially acting as stimulators of a protective response to the fungi. This notion is supported by immunoblotting reactions with a MAb to C. albicans MP65: reactive bands were observed when extracts from C. neoformans acapsular strains were used.
However, we should consider that the observed protection might be due to multiple effects including the stimulation of the innate immune system that could occur through mannose or T-cell-like receptors, at least in the early phase of the immune response. Nevertheless, we can attribute a role to the specific immune response because (i) the non-cross-reactive antigen enolase did not produce a DTH response or affect survival and (ii) adoptive transfer of T cells from CnMP-treated mice induced a DTH response. Thus, MP2 protection may provide early stimulation of a nonspecific immune response with consequent stimulation of specific immunity.
Overall, the evidence for common MP epitopes in C. neoformans and C. albicans suggests at least two considerations: first, that the two fungi synthesize and secrete MP with a potentially similar positive impact on the immune system, and second, that the specific T-cell response could be stimulated, at least in part, by different MPs sharing common epitopes Therefore, identification of the epitopes involved in fungal CMI induction strongly suggests the possibility of using a common protein antigen to elicit a potentially protective adaptive immune response against the fungi.
We thank Jo-Anne Rowe for excellent secretarial and editorial support.
This study was supported by a grant from the National Research Program on AIDS, "Opportunistic Infections and Tuberculosis," contract no. 50D.31, Italy.
- Received 26 April 2002.
- Returned for modification 31 May 2002.
- Accepted 5 September 2002.
- Copyright © 2002 American Society for Microbiology


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26 February, 2011
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I'm going to start by giving you some film stats, about the plot, then go into my review. *WARNING - THIS MAY CONTAIN SPOILERS*
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