President keeps the pressure on Damascus

The anti-Assad line expressed by Mursi in Tehran last week is reiterated at the Arab League in unequivocal terms, reports Dina Ezzat

"Syria, Syria, Syria: You have to do something; it is all up to you and we are here to support you," said President Mohamed Mursi Wednesday morning.

Mursi's statement was made before the opening session of the Arab League Foreign Ministers Council, at the Cairo headquarters of the pan-Arab organisation.

The statement was received with applause from Arab League Secretary-General Nabil El-Arabi and most participating Arab delegations.

Mursi's speech was the first by an Egyptian head of state at the Arab League since ousted President Hosni Mubarak attended the inauguration of a meeting on Iraqi reconciliation seven years ago. Unlike the mild statements Mubarak made then, Mursi's presence and language were uncompromising -- especially on Syria.

"I am telling the ruling regime in Syria that your rule shall not last for long. The Syrian people have said their word, and it is the Syrian people that will have the upper hand," Mursi said.

"I would advise you to refrain from listening to those who are telling you that your rule can persist. I urge you to take the right decision now and not later when it will be too late. You need to part with arrogance and to bow to reality. The Syrian people no longer wants you," he added, addressing -- though he was not present -- Bashar Al-Assad directly.

The president lamented the endless bloodshed suffered by the Syrian people every day, and said that Arab failure to act in support of the Syrian people makes the whole Arab nation responsible for their ordeal.

Mursi's statements came less than a week after harshly criticising the Syrian regime at the opening session of the Non-Aligned Movement summit in Tehran. He went as far as equating the Syrian regime with the Israeli occupation of Palestine.

Mursi's attack elicited a pointed response by a Syrian diplomat who spoke to Al-Ahram Weekly. "Mursi is going the extra mile to appease the Americans. What he is saying is meant to pave the way for wide Arab support for a diplomatic and maybe even military intervention that would be led by the US against Syria," he said.

The Syrian diplomat added that the "attack" is in line with "US directives sent to Cairo through its new Arab ally in Doha". "It is such a shame that Egypt has become a follower of Qatar," he said.

The emir of Qatar is the only head of state to have visited Mursi, mid-August, since he was sworn into office 30 June. Mursi and his Qatari counterpart have met twice since, both on the fringes of an extraordinary summit of the Organisation of the Islamic Conference in Mecca, late August, and in Tehran last week.

A presidential aide who spoke to the Weekly denies that the position adopted by Mursi against Syria is the outcome of Egyptian-Qatari diplomatic coordination. "It is a function of the fact that we really see that this regime [in Damascus] has no chance to stay in power."

Meanwhile, the recently assigned UN-Arab League envoy on Syria, Lakhdar Brahimi, is expected in Cairo in a few days ahead of beginning his mission in the wake of the failure of his predecessor, Kofi Annan.

According to Arab -- including Syrian -- diplomats, Brahimi, a former Algerian foreign minister and an acclaimed international diplomat, has little chance to make a breakthrough in the current crisis in Syria, which started with pro-reform demonstrations in March 2011 and which met a severe military response.

The Syrian opposition, divided as it is, agrees on one thing: Al-Assad has to go. The Syrian regime, however, is not willing to talk about a power transition from Al-Assad and is fighting to survive with some democratic reforms.

Also hindering Brahimi is division within the UN Security Council, with Russia and China still determined to support the Al-Assad regime.

In his speech, Mursi expressed faith in Brahimi.

As Mursi was exiting the Arab League Wednesday, demonstrators gathering not far from the Tahrir Square headquarters of the Arab organisation were shouting: "Down, down with Al-Assad, the butcher of the Syrian people!"

On Tuesday evening, Syrian and sympathetic Egyptian and Arab demonstrators attempted to attack the Syrian embassy in Cairo. Egyptian security forces defended the premises with dozens injured on both sides.

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Deconstructing the third dimension - how 3D culture microenvironments alter cellular cues

Advance Online Publication July 13, 2012 doi: 10.1242/?jcs.079509 July 1, 2012 J Cell Sci 125, 3015-3024. Brendon M. Baker and Christopher S. Chen*

Department of Bioengineering, University of Pennsylvania, 210 South 33rd Street, Philadelphia, PA 19104, USA ?*Author for correspondence (chrischen{at}seas.upenn.edu) Much of our understanding of the biological mechanisms that underlie cellular functions, such as migration, differentiation and force-sensing has been garnered from studying cells cultured on two-dimensional (2D) glass or plastic surfaces. However, more recently the cell biology field has come to appreciate the dissimilarity between these flat surfaces and the topographically complex, three-dimensional (3D) extracellular environments in which cells routinely operate in vivo. This has spurred substantial efforts towards the development of in vitro 3D biomimetic environments and has encouraged much cross-disciplinary work among biologists, material scientists and tissue engineers. As we move towards more-physiological culture systems for studying fundamental cellular processes, it is crucial to define exactly which factors are operative in 3D microenvironments. Thus, the focus of this Commentary will be on identifying and describing the fundamental features of 3D cell culture systems that influence cell structure, adhesion, mechanotransduction and signaling in response to soluble factors, which – in turn – regulate overall cellular function in ways that depart dramatically from traditional 2D culture formats. Additionally, we will describe experimental scenarios in which 3D culture is particularly relevant, highlight recent advances in materials engineering for studying cell biology, and discuss examples where studying cells in a 3D context provided insights that would not have been observed in traditional 2D systems.

Key words Funding

This work was supported in part from grants from the NIH [grant numbers EB00262, EB08396, HL73305, GM74048] and Center for Engineering Cells and Regeneration of the University of Pennsylvania. B.M.B. acknowledges financial support from a Ruth L. Kirschstein National Research Service Award. Deposited in PMC for release after 12 months.

This article is part of a Minifocus on Mechanotransduction. For further reading, please see related articles: ‘Finding the weakest link – exploring integrin-mediated mechanical molecular pathways’ by Pere Roca-Cusachs et al. (J. Cell Sci. 125, 3025-3038). ‘Signalling through mechanical inputs – a coordinated process’ by Huimin Zhang and Michel Labouesse (J. Cell Sci. 125, 3039-3049). ‘United we stand – integrating the actin cytoskeleton and cell–matrix adhesions in cellular mechanotransduction’ by Ulrich S. Schwarz and Margaret L. Gardel (J. Cell Sci. 125, 3051-3060). ‘Mechanosensitive mechanisms in transcriptional regulation’ by Akiko Mammoto et al. (J. Cell Sci. 125, 3061-3073). ‘Molecular force transduction by ion channels – diversity and unifying principles’ by Sergei Sukharev and Frederick Sachs (J. Cell Sci. 125, 3075-3083).

© 2012. Published by The Company of Biologists Ltd

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Controversial new posts

Recent appointments by the Shura Council to the National Council of Human Rights and the state-owned newspapers have renewed concerns over the apparent Islamisation of the country, reports Reem Leila

Members of the Shura Council's general committee met to endorse the final list of members of the Supreme Press Council and to name the new board chairmen of state-owned newspapers on 4 September. The new appointments came just 24 hours after the council had announced the formation of a new National Council of Human Rights (NCHR).

There will be 80 members of the NCHR, 27 of them appointed and forming the council's permanent members. The remaining members will be consultants to the council and will meet with the permanent members once each month. The NCHR's secretary-general will be elected from among the council's appointed members, or chosen from outside the council as stipulated by law.

The appointments made by the Shura Council, the upper house of Egypt's parliament, are the second since the toppling of former president Hosni Mubarak in February 2011. During the meeting of the council's general committee, council speaker Ahmed Fahmi refused nominations made by the council's human rights committee to the NCHR, saying that these should be made "by members of the Shura Council's general committee" instead.

According to Ezzeddin El-Komi, deputy chairman of the Shura Council's Human Rights Committee, the council had agreed to appoint judge Hossam El-Gheriani as head of the NCHR and leftist Abdel-Ghaffar Shokr as deputy. El-Gheriani is known to be an Islamist-leaning figure.

Among the 80 members are four Copts, Georgette Quillini, a lawyer and former MP, Edward Ghaleb, a member of the Coptic Church's Confessional Council, veteran writer Louis Greis, and Ihab El-Kharrat, head of the council's Human Rights Committee.

The council will also include Islamist figures, among them Mohamed Tosson, a prominent lawyer from the Muslim Brotherhood and former chairman of its legislative committee, Abdel-Moneim Abdel-Maqsoud, another Brotherhood lawyer, who defended Khairat El-Shater when the Supreme Electoral Committee excluded the latter from the presidential race, Nader Bakkar, spokesman of the Salafist Nour Party, and Mahmoud Ghozlan, a Brotherhood spokesman who was imprisoned three times in 2002, 2005 and 2007 for opposing the previous regime.

Mohamed El-Beltagui, a prominent leader of the Brotherhood's Freedom and Justice Party (FJP) has also been appointed to the NCHR. El-Beltagui was an MP in 2005, but withdrew from the 2010 parliamentary elections upon the instructions of the Muslim Brotherhood. He is famous for defending the Palestinian cause.

Other figures include Abdallah Badran, head of the Nour Party bloc in the Shura Council, who is known for having criticised former prime minister Kamal El-Ganzouri and members of his government for not attending sessions of parliament. Ahmed Seif El-Islam, secretary-general of the Lawyers Syndicate, is also a member, as are leftist activists Wael Khalil and Mohamed El-Damati, head of the syndicate's Freedoms Committee.

Other nominees include actors Wagdi El-Arabi and Yehia El-Fakharani. El-Arabi declared his affiliation to the Muslim Brotherhood in June 2011.

The NCHR has various topics on its agenda, including social, economic and political issues that have to be addressed in order to grant people their rights and to ensure that there is no discrimination.

Quillini told the Weekly of her happiness at her appointment, saying that in her view there were several controversial issues that needed to be raised at the council. "Despite the recent appointment of Copts to the presidential team, cabinet and NCHR, their representation is still low. The law on unified places of worship should be passed to the People's Assembly for approval," she said.

According to Quillini, there are more Islamists than Copts on the council, but what matters is quality, not quantity. "This is what I intend to prove during my membership. I struggled for Coptic rights when I was an MP, and I will continue fighting until we have reached our goals. We are all equal, and there should not be any discrimination due to differences of religion," she said.

At the same time, writer and novelist Alaa El-Aswani refused his appointment to the council. El-Aswani tweeted: "a free journalist and novelist must not bind himself to any governmental entity in order to be able to criticise the government's performance when necessary without being embarrassed" to do so.

Activist Ahmed Harara, who lost his sight in the course of the 25 January Revolution, also refused his appointment to the council, without giving a reason.

Activist Wael Khalil, who accepted membership of the NCHR, said he would use it to accomplish the goals of the 25 January Revolution. "The council is currently facing serious challenges, among them the reform of the relationship between the government, the police and the people," he said, adding that it would also work to improve the status of women in society, which had been compromised after the revolution.

Activist Mohamed Bahieddin Hassan, head of the Cairo Centre for Human Rights, said that the formation of the new council was proof of the Islamisation of the country and of different organisations and associations.

"The council in its new incarnation will not be independent, and instead it will be controlled by the Muslim Brotherhood. In the past, it was controlled by the now-dismantled National Democratic Party [NDP]," the former ruling party. "It used to appear to be an independent entity, but it was controlled by NDP tycoons. Now there will be no difference: the only difference is that instead of the NDP, it is the FJP that controls everything," Hassan said.

At the same time, the Shura Council also announced new appointments to the Supreme Press Council, including Shura Council head Ahmed Fahmi, eight chairpersons and eight editors-in-chief of state-owned newspapers, and four editors-in-chief of party-affiliated newspapers.

New appointments also included the head of the Press Syndicate, Mamdouh El-Wali, liberal politician Osama El-Ghazali Harb, Ibrahim Hegazi, Mohamed Khuraga and Mohamed Negm.

The new appointments included the head of the General Union of Press, Printing and Publishing Workers, Talaat El-Meneisi. Basiouni Hamdan, Mahmoud Alameddin, legal expert Omar Salem, a former minister of legal affairs, and judge Mohamed Abu Naas.

Several Islamist figures joined the Press Council, such as Salafist Nour Party members Ahmed Khalil and Tarek El-Sahri, and Muslim Brotherhood members Fathi Shehabeddin, head of the Shura Council's Culture Committee, and journalist Qotb El-Arabi.

Osama Ayoub, assistant editor of October magazine, Khaled Salah, editor-in-chief of the newspaper Al-Youm Al-Sabei, Sameh Mahrous, assistant editor of the Al-Gomhuriya newspaper, leftist political figure Wael Qandil, the editor-in-chief of Al-Shorouk newspaper, Naglaa Mahfouz, Ashraf Sadek, Souad Abul-Nasr, and Mohamed El-Abd all also joined the Press Council.

Additional appointments included Geel Party President Nagui El-Shahabi and professor of political science and economics Ayman El-Mahgoub, along with Magdi El-Maasrawi, Mohamed El-Gawadi, Khaled Hassanein, Azza Youssef, Mohamed Hassanein, Ashraf Saber, Abeer Beshr, Hedayat Abdel-Nabi and Mohsen Hassan.

Veteran writer Salah Eissa reportedly objected to the new nominations to the Press Council. "The Shura Council eliminated all those who oppose the Muslim Brotherhood, as well as those who have perspectives on improving the country," he said.

Eissa added that the changes aimed to ensure the control of the Shura Council over the Supreme Press Council, as well as the state-owned newspapers. "They want to implement their own policy without listening to anyone's objections," he said.

In the same session, the Shura Council also announced the appointment of new chairmen of the board of the state-owned newspapers. Chairman of the Press Syndicate Mamdouh El-Wali will be the head of Al-Ahram, instead of Abdel-Fattah El-Gebali. The new chairpersons include Ahmed Sameh at Akhbar Al-Youm, Yehia Zakaria Ghanem at Dar Al-Hilal, Mustafa Abu Zeid at Dar Al-Tahrir, Kamaleddin Mahgoub at Dar Al-Maaref and Shaker Abdel-Fattah at the Middle East News Agency (MENA). Sayed Abdel-Fattah at the National Company of Publications and Mohamed Gamaleddin at Ros El-Youssef have kept their posts.

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Finding the weakest link - exploring integrin-mediated mechanical molecular pathways

Advance Online Publication July 13, 2012 doi: 10.1242/?jcs.095794 July 1, 2012 J Cell Sci 125, 3025-3038. Pere Roca-Cusachs1,*,‡, Thomas Iskratsch2,* and Michael P. Sheetz2,3

1University of Barcelona and Institute for Bioengineering of Catalonia, Barcelona 08028, Spain

2Department of Biological Sciences, Columbia University, New York, NY 10027, USA

3Mechanobiology Institute of Singapore, National University of Singapore, 117411, Singapore?‡Author for correspondence (rocacusachs{at}ub.edu) ?* These authors contributed equally to this work

From the extracellular matrix to the cytoskeleton, a network of molecular links connects cells to their environment. Molecules in this network transmit and detect mechanical forces, which subsequently determine cell behavior and fate. Here, we reconstruct the mechanical pathway followed by these forces. From matrix proteins to actin through integrins and adaptor proteins, we review how forces affect the lifetime of bonds and stretch or alter the conformation of proteins, and how these mechanical changes are converted into biochemical signals in mechanotransduction events. We evaluate which of the proteins in the network can participate in mechanotransduction and which are simply responsible for transmitting forces in a dynamic network. Besides their individual properties, we also analyze how the mechanical responses of a protein are determined by their serial connections from the matrix to actin, their parallel connections in integrin clusters and by the rate at which force is applied to them. All these define mechanical molecular pathways in cells, which are emerging as key regulators of cell function alongside better studied biochemical pathways.

Key words Funding

The work of our laboratory was supported in part by a grant by the Spanish Ministry of Economy and Competitiveness [grant number BFU2011-23111].

This article is part of a Minifocus on Mechanotransduction. For further reading, please see related articles: ‘Deconstructing the third dimension – how 3D culture microenvironments alter cellular cues’ by Brendon M. Baker and Christopher S. Chen (J. Cell Sci. 125, 3015-3024). ‘Signalling through mechanical inputs – a coordinated process’ by Huimin Zhang and Michel Labouesse (J. Cell Sci. 125, 3039-3049). ‘United we stand – integrating the actin cytoskeleton and cell–matrix adhesions in cellular mechanotransduction’ by Ulrich S. Schwarz and Margaret L. Gardel (J. Cell Sci. 125, 3051-3060). ‘Mechanosensitive mechanisms in transcriptional regulation’ by Akiko Mammoto et al. (J. Cell Sci. 125, 3061-3073). ‘Molecular force transduction by ion channels – diversity and unifying principles’ by Sergei Sukharev and Frederick Sachs (J. Cell Sci. 125, 3075-3083).

© 2012. Published by The Company of Biologists Ltd

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Stage demonstration

Gehad Hussein listens to a voice of discontent

As the opposition grows stronger, the monitoring of the current regime becomes more intense. And if a state-owned institution rejects art for political reasons, be sure that the artist will become an effective, critical component of the backbone of the country's opposition -- and thus, influential enough to be censored.

The revolution took place with the aim of getting rid of censorship and not only allow, but also promote freedom of expression, especially in the intellectual and artistic scene. Any setback in that sense is faced with scrutiny and a public scandal on social networks and/or the media.

The theme of this story is a random, ever-changing collection of Egyptians with an undying love for their country and a passion for music. Together, they are called "Mashroua Choral" or "The Choir Project" (TCP). Their artistic director is Salam Youssri, a 30-year-old writer and director. Youssri explains that the aim of TCP is "collective participation. We want to create a safe atmosphere for people to freely express how they feel and what they think."

Being financially independent and dependent on volunteer work, the creation and execution of the choir's projects and songs takes place in workshops and so are free from censorship. Time and place are announced through Facebook and Twitter and anyone can join. "These people learn how to listen to each other and how to stop judging others in a superficial manner," Youssri elaborates. The group sits together and starts brainstorming about what it wants to sing about -- and voil?, a song and sometimes even a whole show is created.

In May 2010, TCP was founded as the Complaints Choir with 22 members who wrote, composed and sang four songs, complaining about everything that was annoying them. The show was portrayed by the media as "singing demonstrations on a stage", as the content of the songs heavily criticised the Mubarak regime. Journalists felt like this was a different way of voicing people's discontent with the political situation -- far from the standard "20 people on the stairs of the Journalists Syndicate" prototype demonstrations at the time. Immediately after the show, State Security threatened the venue Townhouse Gallery, and consequently, it became very hard for TCP to find a venue to perform in, so their songs were mainly published online.

The Choir Project kept rolling, bearing new themes every time, like "Advertising Choir" and "Proverb Choir" and getting invitations to Oman, London, Munich, Berlin, Beirut and Istanbul. Members increased, but the real boost in numbers occurred right after the revolution in their workshop called Utopia where the members sang about the kind of life they wished they had in Egypt.

It seemed like the time of censorship was over, yet, to everyone's disappointment, the mentality of self-censorship still bore its fruits at the core of the state's institution, even those related to art.

As the Cairo Opera House prepared its Ramadan events, it included some revolutionary artists like Rami Essam, Eskenderella and TCP. The performances of the two artists ended with the audiences chanting against the military and army, yet with no direct action or violence. The people working at the Cairo Opera House were not used to this kind of upheaval since they have been working all their lives in a governmental state environment. Hence, the opera's PR manager called Youssri one day before the TCP show, asking him about the content of the songs they were going to perform and informing him that they were not welcome to perform the next day.

According to Youssri, the State Security did not directly call the Opera House and ask them to stop the show, but the people there were used to stopping any opposition before it could get started. Immediately, he wrote a statement about the incident and published it on the Facebook page of the choir. As the social wildfire spread, some journalists started writing about the incident and the story was extremely popular in only a couple of hours. After noticing the heat, the authorities of the Cairo Opera House contacted Youssri again in order to sort out the incident and allowed the Choir Project to perform, after the institution officially apologised for the inconvenience which they branded as a "managerial error."

Shadi El-Husseini, a frequent member of TCP, was not surprised by the incident. "I felt like this would be the normal reaction of a state-owned facility, especially the Hanager Theatre [where the performance took place]. It is just the continuation of the old regime that is still present. It is their loss anyway. But all in all, I believe that this is a positive experience because in the end we were allowed to perform without any censorship."

Youssri further explained: "The move was not specifically against The Choir Project, but rather a fast reaction to what happened in the previous performances with Rami Essam and Eskenderella. As our group is famous for being on the side of the opposition, the Opera House thought it had to act somehow before things got out of hand, which is all a hallucination." He believes that the workers at the Opera House still need to get used to the new things that are going on and to the heat and pressure that social media can trigger.

"In the end, this is our stage, the stage of the people. The Opera House noticed that the self-censorship it had been practicing has no basis," Youssri said.

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The conserved kinase SRPK regulates karyosome formation and spindle microtubule assembly in Drosophila oocytes

Advance Online Publication August 1, 2012 doi: 10.1242/?jcs.107979 In Drosophila oocytes, after the completion of recombination, meiotic chromosomes form a compact cluster called the karyosome within the nucleus, and later assemble spindle microtubules without centrosomes. Although these oocyte specific phenomena are also observed in humans, their molecular basis is not well understood. Here we report essential roles for the conserved kinase SRPK in both karyosome formation and spindle microtubule assembly in oocytes. We have identified a female sterile srpk mutant through a cytological screen for karyosome defects. Unlike most karyosome mutants, the karyosome defect is independent of the meiotic recombination checkpoint. Heterochromatin clustering found within the wild-type karyosome is disrupted in the mutant. Strikingly, a loss of SRPK severely prevents microtubule assembly for acentrosomal spindles in mature oocytes. Subsequently, bi-orientation and segregation of meiotic chromosomes are also defective. Therefore, this study demonstrates new roles of this conserved kinase in two independent meiotic steps specific to oocytes.

This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial Share Alike License, which permits unrestricted non-commercial use, distribution and reproduction in any medium provided that the original work is properly cited and all further distributions of the work or adaptation are subject to the same Creative Commons License terms.

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United we stand - integrating the actin cytoskeleton and cell-matrix adhesions in cellular mechanotransduction

Advance Online Publication July 13, 2012 doi: 10.1242/?jcs.093716 July 1, 2012 J Cell Sci 125, 3051-3060. Ulrich S. Schwarz1,* and Margaret L. Gardel2,*

1BioQuant and Institute for Theoretical Physics, University of Heidelberg, Philosophenweg 19, 69120 Heidelberg, Germany

2Physics Department, James Franck Institute and Institute for Biophysical Dynamics, University of Chicago, Chicago, IL 60637, USA ?*Authors for correspondence (ulrich.schwarz{at}bioquant.uni-heidelberg.de; gardel{at}uchicago.edu) Many essential cellular functions in health and disease are closely linked to the ability of cells to respond to mechanical forces. In the context of cell adhesion to the extracellular matrix, the forces that are generated within the actin cytoskeleton and transmitted through integrin-based focal adhesions are essential for the cellular response to environmental clues, such as the spatial distribution of adhesive ligands or matrix stiffness. Whereas substantial progress has been made in identifying mechanosensitive molecules that can transduce mechanical force into biochemical signals, much less is known about the nature of cytoskeletal force generation and transmission that regulates the magnitude, duration and spatial distribution of forces imposed on these mechanosensitive complexes. By focusing on cell-matrix adhesion to flat elastic substrates, on which traction forces can be measured with high temporal and spatial resolution, we discuss our current understanding of the physical mechanisms that integrate a large range of molecular mechanotransduction events on cellular scales. Physical limits of stability emerge as one important element of the cellular response that complements the structural changes affected by regulatory systems in response to mechanical processes.

Key words Funding

U.S.S. is a member of the Heidelberg cluster of excellence CellNetworks and is supported by the BMBF MechanoSys grant [grant number 0315501C to U.S.S.]. M.L.G. is supported by a NIH Director's Pioneer Award [grant number DP10D00354], Packard Fellowship and Burroughs Wellcome Career Award. Deposited in PMC for release after 12 months.

This article is part of a Minifocus on Mechanotransduction. For further reading, please see related articles: ‘Deconstructing the third dimension – how 3D culture microenvironments alter cellular cues’ by Brendon M. Baker and Christopher S. Chen (J. Cell Sci. 125, 3015-3024). ‘Finding the weakest link – exploring integrin-mediated mechanical molecular pathways’ by Pere Roca-Cusachs et al. (J. Cell Sci. 125, 3025-3038). ‘Signalling through mechanical inputs – a coordinated process’ by Huimin Zhang and Michel Labouesse (J. Cell Sci. 125, 3039-3049). ‘Mechanosensitive mechanisms in transcriptional regulation’ by Akiko Mammoto et al. (J. Cell Sci. 125, 3061-3073). ‘Molecular force transduction by ion channels – diversity and unifying principles’ by Sergei Sukharev and Frederick Sachs (J. Cell Sci. 125, 3075-3083).

© 2012. Published by The Company of Biologists Ltd

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