text:
At its AGM today in Tokyo, 23% of shareholders issued a major climate warning to Mitsubishi UFJ Financial Group (“MUFG”) management by voting in favour of a resolution filed by Kiko Network and three individual shareholders affiliated with Market Forces, Rainforest Action Network, and 350.org Japan (“co-filers”), calling on the bank to adopt and disclose a plan to align its financing and investments with the goals of the Paris Agreement.
In addition to receiving support from investors with trillions of assets under management, such as EOS at Federated Hermes, the resolution proved a catalyst for a major policy shift from MUFG. On May 17, it published the “MUFG Carbon Neutrality Declaration,” and announced its plan to achieve net zero financed emissions by 2050.
As influential international proxy advisor ISS stated in its advice regarding the AGM, ‘we also should state that the bank’s move does not appear unrelated to external pressure: after Kiko Network submitted the proposal in March, MUFG announced its carbon neutrality declaration, so one interpretation of such move is that it was prompted by pressure from the group. That means that Kiko Network’s engagement with MUFG has already successfully borne fruit.”
“Although two major proxy advisors recommended investors vote against the proposal, it is very positive and encouraging that we have nearly one-fourth of investors’ support.
This filing serves as a testament to the impact of investor pressure. Today’s vote is a significant rebuke to management and a clear warning that it must immediately up its game or face even greater pressure,” said Kimiko Hirata, International Director of Kiko Network.
“Faced with the crystallizing physical and financial impacts of the climate crisis, investors are making it plain that MUFG needs to move further and faster.”
Given MUFG’s status as the largest Asian banker of climate chaos, the filers welcomed the May 17 net zero commitment, while noting its numerous shortcomings, including:
- A lack of short and medium term targets required to confirm Paris alignment, stating instead that it “will work to set and disclose an interim milestone for 2030 in FY2022;”
- No explicit commitment to measure and disclose all financed emissions (Scope3);
- No commitment to phase out all coal finance, in contrast with recent demands by the COP26 Presidency;
- No comprehensive commitment to stop financing oil and gas expansion or the destruction of forests and peatlands, both of which are hugely carbon-intensive.
“While MUFG’s new policy has temporarily assured some investors the bank is on the right track, today’s vote shows a high level of dissatisfaction and concern,” said Takayoshi Yokoyama, 350.org Japan Representative. “Even though our proposal was rejected, there were quite a few votes in favor, and as the Corporate Governance Code shows, MUFG must now analyze the reasons for investors’ support and deepen dialogue with its shareholders. MUFG now has the opportunity to close its climate loopholes.”
“Given MUFG’s ranking as the sixth largest banker of fossil fuels in the world and one of the top financiers of rainforest-destroying commodities, its current commitments do not properly address the enormous carbon intensity of its financing and investment portfolio. It is necessary for MUFG to take these supporting votes from its shareholders seriously and take prompt action.” said Toyoyuki Kawakami, Japan Representative of Rainforest Action Network.
“The world is shifting rapidly to clean energy, but MUFG continues to cling onto the past,” said Meg Fukuzawa. “Investors have a responsibility to keep up the pressure and it’s critical for them to apply a razor sharp focus during the next year to ensure MUFG moves from words to genuine action.”


text:
Met Police '˜Flying Squad' make two arrests outside Crystal Peaks
Two men were arrested outside Crystal Peaks in Sheffield in an operation by The Met's '˜Flying Squad'.
Officers in the specialist Metropolitan Police unit, which investigates robberies, swooped on a vehicle as it pulled onto the ‘red’ car park at Crystal Peaks.
The officers arrested two men as part of an investigation into robberies of security vans which carry cash.
The investigation is looking at a number of robberies in London and across the country.
Witnesses who saw officers make the arrests at around 9.30am said a security van was nearby.
The suspects are to be taken to London for questioning later today.
One woman who saw the police operation said: “I was filling up at Sainsbury’s near to Crystal Peaks when I saw a line of cars overtaking all those queuing to get onto the car park.
“I wondered what they were doing and thought how dangerous it was, until I realised they were unmarked police cars when I saw officers in plain clothes make some arrests.”
Crystal Peaks shopping centre is operating as normal today.

domain: news


text:
Nintendo lifts lid on 3DS and new Kid Icarus
The 3DS handheld came to the fore as the culmination of Nintendo's E3 press conference today, the platform holder affirming that '3DS' is the final product name.
The hardware negates the need for 3D glasses, moving the player closer to the action, and there will be lashings of 3D content from day one, we're assured.
The larger screen will be 3.5 inches across, and includes two exterior cameras; allowing users to take 3D pictures. Beyond glasses-free 3D, the 3DS will sport better graphics - good news for third-party developers.
A full motion sensor and gyroscope is included, and the system is compatible with the DSi.
What's more, the 3DS will be capable of playing 3D movies - Nintendo positioning the system as a low-cost way of joining the 3D revolution.
The biggest 3DS game confirmed so far will apparently be Kid Icarus Uprising, a new game being crafted especially to promote the new handheld. An early demo features lots of flying, impressive visuals, and full 3D combat.
Speaking generally, the 3DS will seek-out WiFi spots, downloading new content and software behind the scenes. What's more, there will be no monthly charge for using all this.
Nintencats was confirmed in the briefing, with Kingdom Hearts 3D, Saints Row, DJ Hero 3D, Ninja Gaiden, Street Fighter IV, Professor Layton, Ridge Racer, Metal Gear Solid, and more all on the way. Almost every single big name publisher is involved in some form. We'll have more as we get it.
- Possible dates for the Steam Summer Sale appear online
- Rise Of The Tomb Raider on Xbox 360 doesn't appear in Square Enix's financials for this year
- The Iron Banner returns to Destiny next week with new gear and rewards
- Titanfall developers Respawn won't be at E3 this year
- Destiny: Xur's location and stock for May 28th
- Destiny's Trials Of Osiris declared a success, Bungie bans cheaters
- FIFA 16 will throw women's teams into the mix
- New The Elder Scrolls Online trailer shows off the expansive world of Tamriel
- Batman: Arkham Knight gets a new 7-minute gameplay trailer

domain: entertainment


text:
Five to Nine Cafe
Come and say hello to our Cafe Manager / Barista Jeannie and grab yourself a drink as you drop off your children. Lovely cafe room to sit down and relax, or grab a drink on the run. Open from 8.00 – 10.00am.
- Card making Workshop: Monday 6 August from 9:00am – 11:00am. Frana will be returning to run a card making workshop (small cost to attend)
- Crochet workshop and a knitting workshop (dates yet to be confirmed).
Parent Book Club – details and dates to follow
For more information – see Jeannie in the cafe.
PCG Second Hand Uniform Sale
When: Sunday 8 July
Time: 10.00 – 11.30am
Cash and cheque sales only.
We are always looking for good quality used uniform, particularly polo shirts in size 8 and 10. We are also keen for undergarments and fleeces in size 8.
For more information please click here for the newsletter article.
Order the NEW 2018/2019 Entertainment Book or Entertainment Digital Member ship and you will receive hundreds of valuable offers for everything you love to do, and you will also be supporting Our School PLUS, order now to receive over $200 of bonus Early Bird Offers – hurry these sell out quickly. Click here to order.
Further detailed information can be checked out on the website: http://icept.co.nz/icept/
Job Vacancies Click HERE for current job vacancies.
Before and After School Care
Click here for updated information about before and after school care.
A Day At Clearview
At the start of 2015 we shared a video that Tony Stewart created to show the amazing things that happen at Clearview each day. To check it out, click on this link: http://youtu.be/j1HzyfHMwrU
Published on Tuesday, February 4th, 2014, under Notices

domain: hobby


text:
Nowhere I could find the real pictures of the Ryabinins great Russian poets, epic story tellers. This is why it is shown here a piece of an artist’s imagination of old Russian story tellers. To attune your ear to the Russian ancient melody it is included also the performance of folk Russian group “Belorybitsa”, song Рябина (Ryabina)
On December 3, 1871 the newspaper “Russian World” wrote: “There is no doubt that to listen to bylinas from lips of the story teller (rapsoda) is a case, rare and rather interesting, drew attention to enormous public who stood on foot in the hall because of lack of seats. In front of the listeners at a table in an armchair, having put his both hands on arm rests, the old man of sturdy build, with a short growth of grey beard and yellow hair sat. It was Trofim Grigorievich. At first sight he seemed bent over in the way he walked, his outward appearance was one which impresses with strength and endurance. Dressed in sibirku, his face with deeply set eyes and all his outlooks show a respectable and serious kind.
With his beautiful, profound, live, cheerful voice the tempo of which was at times faster, at other times the harmony was broken reminding us of something age-old, forgotten by our generations… “Ilja Muromets and Nightingale the Robber” started approximately as follows:
Whether from here or there perhaps from the city of Murom,
Oh hey, maybe from the village of Karacharovo,
A good fellow was traveling on a long, long road,
He stopped in the early morning near Murom,
And by noon he kept on his way to the capital city of Kiev,
Oh hey, meanwhile he approached a nice city of Chernigov,
Whether at that city of Chernigov either close or far,
He was put on force trial by something pitch-dark,
And hey, pitch-dark was a black raven…
13 texts had been written down, among them: “Ilja and Idolishche”, “Ilja and Guelder-roses-tsars”, “Ilja in Quarrel with Prince Vladimir”, “Dobrynja and Dragon”, “Dobrynja and Marinka, “Volga and Mikula”, ” The Good Fellow and a Thin Wife” etc.Characters of Russian folklore
Listening to the bylinas we are plunging into the special epic world, both fantastic and real. After Trofim Grigorievich Ryabinin’s performance he was handed over a monetary compensation of the amount being 100 silver roubles, and Grand Duke Konstantin Nikolaevich had awarded the singer really with a “princely” gift the silver cup executed in old Russian style, Silver medal, 1872 -official seal.
The National Archives read:
“The Sovereign the Emperor in Response to the report of the Minister of the Internal Affairs, Its desire of Imperial Highness of the Sovereign of Grand Duke Konstantin Nikolaevich, about rewarding the well known rapsoda and the story teller of Russian bylinas, the peasant of Olonetsky province Trofim Ryabinin, is more graciously deigned to welcome on the 18-th day of February to Ryabinin the Silver Medal with an inscription “for useful” for carrying on a neck on to a Stanislavsky tape.” In Match 1872 the Olonetsky governor G. Grigoriev had called T. Ryabinin to Petrozavodsk and personally handed the award over to him.
Trofim Grigorievich Ryabinin was born on April 15, 1801 (27-th April-new style) in v. Dmitrovsky (Gamitsky-the present name). Very early he had lost his parents and had been brought up in the “country world”, “world” means godfathers, matchmakers, godmothers, aunts, uncles, “daleekieedaleeekiee”.
In 1820 he married to Darya Pankratovna Klopovaja, came to her parents’ house as a “primak”. Later he had built up his own house, the couple had 16 children.Черный баран (Black Ram)
On February 14, 1885 Trofim Ryabinin had died. According to his son Ivan, “very much driven was: went, went and had died a safe death”. He had left imperceptibly in a non-existence, in immortality. He had been buried in Kizhi at the church yard, about Pokrovsk Churches.
His son Ivan Trofimovich Ryabinin became a successor of a poetic talent. He adopted the basic part of repertoire, 15 bylinas, almost literally repeated the texts of his father. The epic songs were performed in front of grateful listeners of St. Petersburg, Moscow, Odessa, Sofia, Belgrade, Vienna, Prague, Warsaw.
Anton Arensky (1861-1906) was inspired to compose Fantasia on Themes by I.T. Ryabinin, Op.48 for piano and orchestra, and one bylina from Novgorod had been the inspiration behind Rimsky-Korsakov’s opera “Sadko”.
In this Ryabinin Fantasia, perhaps modeled on comparable works by Liszt, Arensky chose two themes borrowed from the collection of Trofim’s son Ivan (1845-1910) The first theme (Andante sostenuto) is a gentle, lyrical melody in E minor presented first by the piano and then by the orchestra with embellishments from the soloist. The second theme is a march-like theme in D minor (Allegretto) with a “scotch snap” rhythm, the florid style is often used, the sounds of the gusli (zither) so closely associated with the Russian bard tradition.
Nowadays 208 years later we are celebrating the date of the birth of the first great Russian poet among the best story tellers Trofim Grigorievich Ryabinin, the talented peasant fisherman from the northern Onega region of Russia, the founder of the dynasty of bards who kept the oral tradition of the ancient bylinas (epic songs) alive.

domain: artistic


text: A city-ordered investigation into a feud between Ferndale police and the county prosecutor will determine what the City Council’s next move is on the issue, officials said. “We would like to get back to a situation where our local police and the prosecutor’s office are working closely together and in tandem,” said Mayor Craig Covey. In most cities, the police department is subordinate to the city manager’s office. In Ferndale, however, all department heads are appointed directly by the City Council which oversees them. Covey said officials won’t know whether disciplinary action against any officers may be warranted until the investigation is completed. “At this point, we are not looking for disciplinary action,” Covey said. “We are looking to solve the problem.” The City Council on Wednesday voted 4-0 to hire the city labor attorney’s firm to look into the dispute between police and Oakland County Prosecutor Jessica Cooper and her office. Because of issues arising from the conflict with Ferndale police, Cooper’s office requires all Ferndale police requests for warrants to be presented in person in Pontiac. Cooper no longer allows her office to communicate with city police by phone or e-mail when they seek warrants. Police Chief Michael Kitchen has said appearing in person has so far cost more than $500 for time and travel for police officers. Problems between police and Cooper’s office began last fall when Ferndale police publicly criticized prosecutors for failing to issue criminal charges against a man they say pistol-whipped a teen in the Ferndale High School parking lot. The friction abated after police apologized to the prosecutor, Kitchen has said. But problems reignited last month when police released tapes of phone calls with prosecutors in a domestic violence investigation where prosecutors refused to issue a criminal warrant. Ferndale and a number of other police departments record phone calls. Police released the tapes to a TV news station which requested the recordings under a Freedom of Information Act request. Kitchen said the FOIA release was first reviewed by the city attorney. He also has said the family in the domestic assault case had contacted the TV station about the issue, not police. Cooper this week said her office would not “be extorted into writing (criminal) warrants that are not appropriate.” She also noted she has not had problems with the other 46 police departments in the county. She further faulted Ferndale police for releasing taped phone calls that she said are exempt from FOIA requests. Though the City Council voted to begin its investigation into the issue, Councilman Mike Lennon was not at the meeting and objects to spending money for such an inquiry. “I think it’s a waste of money,” said Lennon, a retired Ferndale police officer. The city pays $105 per hour to the labor attorney whose firm will investigate the case, according to city records. “It’s going to cost us quite a bit,” Lennon said. “There are a lot of issues here, but the ones who are suffering are Ferndale residents who have to pay to send officers up to Pontiac every time they need a warrant. If I had my way, we’d put the people involved around a table and settle it.” Kitchen maintains that Cooper is unwilling to meet with him. “She still thinks we called the media and that absolutely didn’t happen,” Kitchen said.


text:
SEN3: An exciting new genre-hopping guitar trio have emerged from London’s melting pot. The band create a compelling wide-screen soundworld that’s rich in melody and emotive harmonies, contemporary beat-culture and heavy riffs. Imagine a combo of Pink Floyd’s ethereal epic-ness, Jimi Hendrix’s psychedelic soul-funk, a touch of J-Dilla/Madlib beat conducting with a twist of Led Zepellin riffs and you’re half way there.
With credits as in-demand sidemen for David Gilmour, Jamie Woon, Jessie Ware, Khadi Tatham and PBUG, the band stretched out in early 2017, releasing their debut album The Drop to rave reviews and plenty of attention from Jazz FM, BBC Radio, Ronnie Scott’s, Jazzwise Magazine and Dublab. Their second studio album Volume II is out now and available on bandcamp and all streaming platforms. Recent gigs include Love Supreme festival, Cheltenham Jazz Festival, Ronnie Scotts Jazz Club (Supporting Sly and Robbie). SEN3 are ones to watch
“Love this new Project” Chris Philips, Jazz FM
“SEN3 are in possession of a seriously unique and energetic sound – loving theses guys.” Tony Minvielle, Jazz FM
“I fully support this exciting new project and these guys will blow up for sure!” Rebecca Vasmant, BBC Radio Scotland
“SEN3 are killing it, as a live band they are a force to be reckoned with” Nick Lewis, Ronnie Scott’s


text:
The complex legal wrangling between the owner of the Versace Mansion on Miami Beach, his lenders and restaurateur Barton Weiss has taken its toll, contributing to a 25 percent reduction in the asking price for the property to $75 million.
Reading between the lines, it’s easy to understand the motivation behind the change from $100 million, which the property’s marketing team Jill Hertzberg and Jill Eber announced Wednesday.
“When the Casa Casuarina hit the market, it generated worldwide interest from prospective buyers, but there was hesitation because there was a tenant occupying the property,” Hertzberg said in a news release. “Now with a non-issue, combined with a motivated seller, we look forward to showcasing this one-of-a-kind mansion and finding the right buyer for it.”
Actually, Weiss filed a lawsuit in early May complaining that property owner Peter Loftin did not fulfill his end of the bargain when both signed on to locate The Villa by Barton G restaurant at the mansion. That lawsuit is still active.
Loftin is also now back in bankruptcy court because of the property’s link to Ponzi schemer Scott Rothstein, who made a $1 million investment in the property in exchange for a minority position in Loftin’s company.
The bankruptcy trustee overseeing Rothstein’s defunct law firm had dropped his claim on the mansion, but changed his mind once the property was marketed for sale at $100 million.
The Nakash family, of Jordache jeans fame, paid $14 million for part of the debt on the property and they are fighting to take control.
In the center of the legal tug of war is the mansion: the 10-bedroom, 11-bathroom, gated home includes a 54-foot-long, mosaic pool lined in 24-karat gold; numerous frescoes and statues; arched doorways; and an open air courtyard.
Built in 1930 by Alden Freeman, Gianni Versace bought the property in the early ‘90s. He invested more than $30 million into the property including an expansion. He was murdered in 1997 in front of the mansion by serial killer Andrew Cunanan.
Loftin kept all of Versace’s design elements after buying the property in 2000.

domain: economics


text:
Justin Timberlake is almost finished writing his second album.
The singer has penned over 15 potential tracks with longtime friend Matt Morris and is planning to hit the studio before the summer is out.
"Writing with Justin is one of my favourite things," Morris told MTV.com. "He's got incredible instincts, unmatched by anyone I've worked with."
The subject matter of the new songs is said to be so personal that discussing them "should be reserved for close friends and therapists".

domain: celebrity


text:
A panel of US House of Representatives established earlier on the present time that four important technology corporations are abusing their market vitality to murder opponents and drive minute companies to their knees within the name of earnings.
In accordance with the antitrust subcommittee of the Judiciary Committee, Google, Apple, Amazon, and Facebook ought to not retain an eye on and compete within the identical industry. The panel suggested these corporations ought to be restructured but didn’t imprint whether a particular firm ought to be broken up, as per the Reuters fable.
The elephantine fable is 449 pages lengthy and suggests noteworthy changes to antitrust law and described “dozens of instances where corporations misused their vitality”. It also printed how corporations did all the pieces they would possibly well to dominate over competitors in uncover to retain an eye on giant portions of the internet.
One instance is Facebook’s acquisition of Instagram wait on in 2012, when Label Zuckerberg, Facebook CEO, illustrious the picture-sharing platform is constructing a aggressive network that will perhaps furthermore very smartly be “very disruptive to us”, the fable reads.
Changes suggested by the committee consist of stopping Google to both gallop the auctions for online ad space and dangle half in these auctions. One other recommendation is for Amazon to end running on markets where it also competes with diversified retail outlets, but the fable also entreated US Congress to behave on allowing antitrust enforcers more freedom to end purchases of potential opponents.
The fable used to be printed by a committee led by Democratic Congressman David Cicilline. Reuters illustrious that if the presidential elections are gained by Joe Biden, the Democratic majority within the House would possibly well retain pressuring Congress to behave on the findings and introduce a revamped antitrust law.


text:
Hello, friends! I have a long list of posts that I'm planning to write, once I have some time. I've been busy having fun, yes, but I've also been hard at work on the second draft of my novel, a process which has been humbling, exciting, and - in rare moments - transcendent. It's a good thing I love revision, because my book needs a lot of it!
In the meantime, I have a short essay published on Elephant Journal this morning. It's about Morning Glory Yoga, the studio where I spent so much time in Nacogdoches, but it's also about friendship, the idea of home, and the uncomfortable reality of change. If you read it, let me know what you think, and feel free to share the link with every single person you know. Not because this is a paying gig (it's not, unfortunately), but because writing is so often a solitary experience, and the opportunity to share my work is always thrilling. Plus I'm really proud of the title, as puns are the best.
Hope you're having a good day, wherever you are. I'll be back soon. <3

domain: literature


text:
German police said they had arrested an arms dealer who sold the Munich shooter a gun and nearly 300 bullets, said on its site newspaper “Bild”, quoted by Tass.
According to the newspaper the 31-year-old man from Marburg, Hesse, was under surveillance in recent weeks. Five days before the attack on July 22, 18-year-old German of Iranian origin David Sonboli Ali bought his Glock 17 handgun and ammunition on the Internet using the program Dark Net to conceal his tracks.
Investigators made contact with the suspect and ordered a Glock 17 and bullets totaling € 8,000. When transferring the weapon and ammunition in Marburg the man was arrested. He could face up to 10 years in prison for violating the law for purchasing and storing weapons.
On July 22 in a shopping center in Munich, 18-year-old Sonboli shot nine people and then killed himself. Injured were 35 people. Thereafter was clarified that the young man was interested in massacres and suffered from unstable psyche with panic attacks and depression.

domain: law


text:
Mario Kart 8 Deluxe New Courses. For a gallery of images related to the original mario kart 8 installment, see gallery:mario kart 8. The second wave of new dlc tracks for mario kart 8 deluxe has only just.
The second wave of the mario kart 8 deluxe dlc is arriving on august 4, 2022. Mario kart 8 deluxe is a racing game in the mario kart series for the nintendo switch which was first released on april 28, 2017. All of the included courses can be found below:
Mario Kart 8 Deluxe (Stylized As Mariokart 8 Deluxe And Abbreviated To Mk8Dx) Is A Mario Kart Racing Game Exclusive To The Nintendo Switch.
Mario kart stadium • water park • sweet sweet canyon • thwomp ruins: 200cc has the speed cranked so high that the ai can barely handle it. It amounted to a glorified reskin at the time, bundling all the.
Mario Kart 8 Deluxe Players Will Once Again Find Two New Cups To Jump Into, With A Total Of Four Tracks For Each Cup Arriving In Wave 2.
The game is the first mario game overall to be released for the nintendo switch. Mario kart 8 deluxe players have a wealth of content to look forward to with the booster course pass dlc which is set to add a whopping 32 courses to the game over an extended period of time. Some of these tracks included toad circuit (3ds), choco.
It Is An Enhanced Port Of Mario Kart 8 For The Wii U.it Has Additional Features Such As Several New Characters, Many New Dlc Courses (Added As Of 2022.
This is a gallery of images for the game mario kart 8 deluxe and its downloadable content, the booster course pass. Mario kart 8 deluxe is an enhanced port of the wii u game, including new characters, a revamped battle mode, and the ability to hold two items at once, not seen since double dash!!.also inklings. Yesterday's announcement revealed which new tracks will be.
Courses New Courses Mushroom Cup:
If you want to check out the new courses, you can grab the mario kart 8 dlc for £22.49 from. Depending on the track played, you can see the ai smash into the walls or go flying off the track and into a bottomless pit. It is the enhanced port (or reboot) version of mario kart 8 and was released worldwide on april 28, 2017.
Battle On New Courses, Like Urchin Underpass And Battle Stadium, Or Returning Ones, Such As Gcn Luigi's Mansion And Snes Battle Course 1.
Mario kart 8 deluxe has new options to help out rookie racers, like smart steering. For a gallery of images related to the original mario kart 8 installment, see gallery:mario kart 8. It was released in south korea on december 15, due to the nintendo switch being released on december 1 there.


text:
Chimpanzees are among the closest living relatives to humans and, as such, provide a crucial comparative model for investigating primate brain evolution. In recent years, human brain mapping has strongly benefited from enhanced computational models and image processing pipelines that could also improve data analyses in animals by using species-specific templates. In this study, we use structural MRI data from the National Chimpanzee Brain Resource (NCBR) to develop the chimpanzee brain reference template Juna.Chimp for spatial registration and the macro-anatomical brain parcellation Davi130 for standardized whole-brain analysis. Additionally, we introduce a ready-to-use image processing pipeline built upon the CAT12 toolbox in SPM12, implementing a standard human image preprocessing framework in chimpanzees. Applying this approach to data from 194 subjects, we find strong evidence for human-like age-related gray matter atrophy in multiple regions of the chimpanzee brain, as well as, a general rightward asymmetry in brain regions.
Chimpanzees (Pan troglodytes) along with bonobos (Pan paniscus) represent the closest extant relatives of humans sharing a common ancestor approximately 7–8 million years ago (Langergraber et al., 2012). Experimental and observational studies, in both the field and in captivity, have documented a range of cognitive abilities that are shared with humans such as tool use and manufacturing (Shumaker et al., 2011), symbolic thought (de and Frans, 1996), mirror self-recognition (Anderson and Gallup, 2015; Hecht et al., 2017) and some basic elements of language (Savage-Rumbaugh, 1986; Savage-Rumbaugh and Lewin, 1994; Tomasello and Call, 1997) like conceptual metaphorical mapping (Dahl and Adachi, 2013). This cognitive complexity together with similar neuroanatomical features (Zilles et al., 1989; Rilling and Insel, 1999; Gómez-Robles et al., 2013; Hopkins et al., 2014; Hopkins et al., 2017) and genetic proximity (Waterson et al., 2005) renders these species unique among non-human primates to study the evolutional origins of the human condition. In view of evolutionary neurobiology, the relatively recent divergence between humans and chimpanzees explains the striking similarities in major gyri and sulci, despite profound differences in overall brain size. Numerous studies using magnetic resonance imaging (MRI) have compared relative brain size, shape, and gyrification in humans and chimpanzees (Zilles et al., 1989; Rilling and Insel, 1999; Gómez-Robles et al., 2013; Hopkins et al., 2014; Hopkins et al., 2017).
Previous studies of brain aging in chimpanzees have reported minimal indications of atrophy (Herndon et al., 1999; Sherwood et al., 2011; Chen et al., 2013; Autrey et al., 2014). Nevertheless, Edler et al., 2017 recently found that brains of older chimpanzees’ exhibit both neurofibrillary tangles and amyloid plaques, the classical features of Alzheimer’s disease (AD). Neurodegeneration in the aging human brain includes marked atrophy in frontal and temporal lobes and decline in glucose metabolism even in the absence of detectable amyloid beta deposition, which increases the likelihood of cognitive decline and development of AD (Jagust, 2018). Given the strong association of brain atrophy and amyloid beta in humans, this phenomenon requires further investigation in chimpanzees.
Cortical asymmetry is a prominent feature of brain organization in many primate species (Hopkins et al., 2015) and was recently shown in humans in a large-scale ENIGMA (Enhancing Neuroimaging Genetics through Meta-Analysis) study (Kong et al., 2018). For chimpanzees, various studies have reported population-level asymmetries in different parts of the brain associated with higher order cognitive functions like tool-use (Freeman et al., 2004; Hopkins et al., 2008; Hopkins et al., 2017; Hopkins and Nir, 2010; Lyn et al., 2011; Bogart et al., 2012; Gilissen and Hopkins, 2013) but these results are difficult to compare within and across species, due to the lack of standardized registration and parcellation techniques as found for humans.
To date, there is no common reference space for the chimpanzee brain available to reliably associate and quantitatively compare neuro-anatomical evidence, nor is there a standardized image processing protocol for T1-weighted (T1w) brain images from chimpanzees that matches human imaging standards. With the introduction of voxel-based morphometry (Ashburner and Friston, 2000) and the ICBM (international consortium of brain mapping) standard human reference brain templates almost two decades ago (Mazziotta et al., 2001), MRI analyses became directly comparable and generally reproducible. In this study, we adapt state-of-the-art MRI (magnetic resonance imaging) processing methods to assess brain aging and cortical asymmetry in the chimpanzee brain. To make this possible, we rely on the largest openly available resource of chimpanzee MRI data: the National Chimpanzee Brain Resource (NCBR, http://www.chimpanzeebrain.org/), including in vivo MRI images of 223 subjects from 9 to 54 years of age (Mean age = 26.9 ± 10.2 years). The aim of this study is the creation of a chimpanzee template permitting automated and reproducible image registration, normalization, statistical analysis, and visualization to systematically investigate brain aging and hemispheric asymmetry in chimpanzees.
Initially, we created the population-based Juna.Chimp (Forschungszentrum Juelich - University Jena) T1-template, tissue probability maps (TPM) for tissue classification and a non-linear spatial registration ‘Shooting’ templates (Figure 1) in an iterative fashion at 1 mm spatial resolution. The preprocessing pipeline and templates creation were established using the freely available Statistical Parametric Mapping (SPM12 v7487, http://www.fil.ion.ucl.ac.uk/spm/) software and Computational Anatomy Toolbox (CAT12 r1704 http://www.neuro.uni-jena.de/cat/). Juna.Chimp templates, the Davi130 parcellation, gray matter (GM) masks utilized, as well as statistical maps from our analysis are interactively accessible and downloadable via the Juna.Chimp web viewer (http://junachimp.inm7.de/).
To enable more direct comparison to previous research, we manually created the Davi130 parcellation (by R.D. and S.V.), a whole brain macroanatomical annotation based on the Juna T1 template (Figure 2). The delineation of regions within the cortex was determined by following major gyri and sulci, whereby, large regions were arbitrarily split into two to three sub-regions of approximate equal size even though histological studies show that micro-anatomical borders between brain regions are rarely situated at the fundus (Sherwood et al., 2003; Schenker et al., 2010; Spocter et al., 2010; Amunts and Zilles, 2015). This process yielded 65 regions per hemisphere for a total of 130 regions for the Davi130 macro-anatomical manual parcellation (Figure 2 and Figure 2—source data 1).
Following successful CAT12 preprocessing, rigorous quality control (QC) was employed to identify individual MRI scans suitable for statistical analysis of brain aging and hemispheric asymmetry in chimpanzees. Our final sample consists of 194 chimpanzees including 130 females with an age range of 9–54 years and a mean age of 26.3 ± 9.9 years (Figure 3A). The linear regression model with GM fraction of total intracranial volume as the dependent variable and age, scanner field strength, sex, and rearing environment revealed a significant negative association between age and GM (p<0.0001) demonstrating age-related decline in overall GM density (Figure 3B). Both sex (p=0.004) and scanner field strength (p<0.0001) showed a significant effect on total GM volume. Therefore, the sample was split into male and female subjects and into 1.5T and 3T scanner, whereby, all sub-samples showed a significant age effect on GM (male: R2 = 0.17, p=0.0004; female R2 = 0.13, p<0.0001, 1.5T: R2 = 0.19, p<0.0001; 3T: R2 = 0.09, p=0.004). There were no significant sex differences of GM decline (p=0.3). The same analysis was conducted on a matched human sample from the IXI dataset (Figure 3C; https://brain-development.org/ixi-dataset/). The human sample was matched based on age, sex, and scanner field strength (n = 194, 128 females, 20–78 y/o, mean = 39.4 ± 14.0). As life span and aging processes are different between species, the human sample was matched to chimpanzees roughly by using a factor of 1.5* for age. A significant age-related decline in overall GM (p<0.0001) as well as a significant sex effect (p<0.0001) was also found in the human sample (Figure 3D). Similar to the chimpanzee sample, both males and female subjects show a significant age effect on total GM (male: R2 = 0.58, p<0.0001; female: R2 = 0.61, p<0.0001) but with no significant sex differences on GM decline (p=0.8). Although both species present a significant age-related GM decline, humans show a higher negative correlation between age and GM (chimpanzee: R2 = 0.12; human: R2 = 0.55) with less variance as compared to chimpanzees.
Region-based morphometry analysis was applied to test for local effect of age on GM. Linear regression analyses identified 55 of 130 brain regions in the Davi130 parcellation across both hemispheres that were significantly associated with age after family-wise error (FWE) correction for multiple testing (Figure 4 and Figure 4—source data 1). Specifically, GM decline with age was found bilaterally in the superior frontal gyrus (SFG), posterior middle frontal gyrus (pMFG), posterior inferior frontal gyrus (pIFG), lateral orbitofrontal cortex (lOFC), middle and inferior precentral gyrus (PrCG), cingulate gyrus (ACC, MCC, PCC), posterior superior temporal gyrus (pSTG), anterior middle temporal gyrus (aMTG), precuneus (PCun), and lingual gyrus (LG) as well as unilaterally in the right anterior insula (aIns) and middle inferior frontal gyrus (mIFG), in addition to the left superior precentral gyrus (sPrCG), anterior transverse temporal gyrus (aTTG), posterior transverse temporal gyrus (pTTG), paracentral lobule (PCL) and the area around the calcarine sulcus (Calc) within the cerebral cortex. Subcortically, age-related GM decline was found in the bilateral putamen (Pu), caudate nucleus (CN), and the nucleus accumbens (NA), as well as in the superior cerebellum (CerVI, CerIV, CerVA, Cer VB, CerIV and right CrusII). Finally, to test for more fine grained effects of aging independently of our macroanatomical parcellation, the same sample was analyzed with VBM revealing additional clusters of GM that are significantly affected by age in chimpanzees (Figure 5) after FWE correction using threshold-free cluster enhancement (TFCE) (Smith and Nichols, 2009). On top of the regions identified by region-wise morphometry, we found extensive voxel-wise effects throughout the orbitofrontal cortex (OFC), inferior temporal gyrus (ITG), transverse temporal gyrus (TTG), frontal operculum (FOP), parietal operculum (POP), postcentral gyrus (PoCG), supramarginal gyrus (SMG), angular gyrus (AnG), and in parts of the superior parietal lobule (SPL), superior occipital gyrus (sOG), and in inferior parts of the cerebellum.
Hemispheric asymmetry of the chimpanzee brain was assessed for each cortical Davi130 region with a total of 68% (44/65) exhibiting significant cortical asymmetry after FWE correction (Figure 6 and Figure 6—source data 1). The majority of regions were found with greater GM volume in the right hemisphere (n = 32) as compared to the left (n = 12). In the left hemisphere, we found more GM in the SFG, pMFG, insula, anterior TTG, and PCun within the cortex. Rightward cortical asymmetry was located in the anterior MFG, middle and posterior IFG, medial OFC, cingulate gyrus, amygdala, STG, MTG, posterior TTG, anterior and posterior fusiform gyrus (FFG), FOP, POP, middle PrCG, middle and inferior PoCG, SMG, AnG, Calc, as well as the middle occipital gyrus. Within the basal ganglia, leftward GM asymmetry was observed in the Pu, nucleus accumbens (NA), basal forebrain nucleus (BF), and globus pallidus (GP), while, rightward asymmetry in the caudate nucleus (CN) and thalamus (Th). The cerebellum exclusively showed rightward GM asymmetry in the posterior cerebellar lobe (CerIX, CerVIII, CrusI, CrusII, CerVI). The hemispheric asymmetry did not show a decipherable pattern.
As a common reference space for the analysis of chimpanzee brain data, we created the Juna.Chimp template, constructed from a large heterogeneous sample of T1w MRI’s from the NCBR. The Juna.Chimp template includes a reference T1-template, along with probability maps of brain and head tissues accompanied by a Geodesic Shooting template for the publicly available SPM12/CAT12 preprocessing pipeline to efficiently segment and accurately spatially normalize individual chimpanzee T1w images. The T1-template and TPM can also be used as the target for image registration with other popular software packages, such as FSL (https://fsl.fmrib.ox.ac.uk/fsl) or ANTs (http://stnava.github.io/ANTs/). Furthermore, our processing pipeline and templates can be utilized for data-driven approaches to create connectivity-based and structural covariance parcellations of the chimpanzee brain (Alexander-Bloch et al., 2013; Eickhoff et al., 2015).
Additionally, we provide the manually segmented, macro-anatomical Davi130 whole-brain parcellation comprising 130 cortical, sub-cortical and cerebellar brain regions, which enables systematic extraction of volumes-of-interest from chimpanzee MRI data. The image processing pipeline and Davi130 parcellation were used to investigate ageing and interhemispheric asymmetry in the chimpanzee brain. The Davi130 parcellation was realized by utilizing macroscopic gyral and sulcal features such as peaks, fundi, and bends to represent the chimpanzee brain in a reduced dimensional space based on macro-anatomical landmarks. Despite the evidence that true microanatomical borders between brain areas rarely coincide with macro-anatomical patterns (Amunts and Zilles, 2015), macroscopic brain parcellations like Desikan-Killany human atlas (Desikan et al., 2006) have successfully been utilized in many studies furthering our understanding of brain structure, function, and disease (Kong et al., 2020; van den Heuvel et al., 2020). The Davi130 parcellation of the chimpanzee brain serves two main purposes. First, regions in Juna.Chimp template space enable increased interpretability and reproducibility of morphometric analyses and comparability between studies, even retrospectively. Second, our manual subdivision reduces the statistical problem of multiple testing for mass univariate approaches like VBM to uncover subtle brain - behavior relationships. Furthermore, our macroscopic parcellation mitigates the curse of dimensionality for multivariate machine learning methods to be applied to the relatively small samples like the NCBR.
We found clear evidence of global and local GM decline in the aging chimpanzee brain even though previous research into age-related changes in chimpanzee brain organization has shown little to no effect (Herndon et al., 1999; Sherwood et al., 2011; Chen et al., 2013; Autrey et al., 2014). (Herndon et al., 1999; Sherwood et al., 2011; Chen et al., 2013; Autrey et al., 2014). This can be attributed on the one hand to the larger number of MRI scans available via the NCBR including 30% of older subjects with 55 individuals over 30 and 12 over 45 years of age, which is crucial for modelling the effect of aging (Chen et al., 2013; Autrey et al., 2014). On the other hand, state-of-the-art image processing enabled the creation of the species-specific Juna.Chimp templates, which largely improves tissue segmentation and registration accuracy (Ashburner and Friston, 2000). Non-linear registration was also improved by the large heterogeneous sample utilized for the creation of the templates encompassing a representative amount of inter-individual variation. We used the well-established structural brain imaging toolbox CAT12 to build a reusable chimpanzee preprocessing pipeline catered towards analyzing local tissue-specific anatomical variations as measured with T1w MRI. The Davi130-based region-wise and the voxel-wise morphometry analysis consistently showed localized GM decline in lateral frontal cortex, lOFC, precentral gyrus, cingulate gyrus, PCun, medial parietal and occipital cortex, the basal ganglia, and superior cerebellum. The VBM approach additionally produced evidence for age effects in bilateral mOFC, PoCG, inferior temporal regions, inferior and superior lateral parietal cortex, sOG, and throughout the cerebellum. These additional effects can be expected, as VBM is more sensitive to GM changes due to aging (Kennedy et al., 2009). The multiple brain regions revealing GM decline reported here in both approaches have also been shown to exhibit GM atrophy during healthy aging in humans (Good et al., 2001b; Kennedy et al., 2009; Crivello et al., 2014; Minkova et al., 2017). Additionally, there was no significant difference in the age-related decline between humans and chimpanzee (Figure 3), even though a larger negative correlation with less variance was found in the matched human sample, which demonstrates a commonality in the healthy aging process of chimpanzees that was thought to be specific to humans. In general, GM atrophy in chimpanzees occurs across the entire cortex, sub-cortical regions, and cerebellum, however, certain local areas decline at a relative extended rate within the frontal, temporal, and parietal lobes (Fjell et al., 2014). Several Davi130 regions within the frontal lobe (SFG, MFG, IFG, and lOFC) have been previously reported in corresponding human loci in relation to GM volume decline due to aging (Kennedy et al., 2009; Crivello et al., 2014; Fjell et al., 2014; Minkova et al., 2017). Furthermore, aging effects in temporal (STG) and parietal (PoCG, AnG, PCun) regions in chimpanzees have additionally been revealed in analogous human areas (Good et al., 2001b; Kennedy et al., 2009; Crivello et al., 2014; Fjell et al., 2014; Minkova et al., 2017). The same is true for the superior occipital gyrus and caudate nucleus (Crivello et al., 2014). Similar age-related total GM decline along with presentation in homologous brain areas suggests common underlying neurophysiological processes in humans and chimpanzees due to shared primate evolution.
Very recently, it has been shown that stress hormone levels increase with age in chimpanzees, a process previously thought to only occur in humans, which can cause GM volume decline (Emery Thompson et al., 2020). This further strengthens the argument that age-related GM decline is also shared by humans closest relative, the chimpanzee. Furthermore, Edler et al., 2017 found Alzheimer’s disease-like accumulation of amyloid beta plaques and neurofibrillary tangles located predominantly in prefrontal and temporal cortices in a sample of elderly chimpanzees between 37 and 62 years of age. As the aggregation of these proteins is associated with localized neuronal loss and cortical atrophy in humans (La Joie et al., 2012; Lladó et al., 2018), the age-related decline in GM volume shown here is well in line with the findings by Jagust, 2016 associating GM atrophy with amyloid beta. These findings provide a biological mechanism for accelerated GM decrease in prefrontal, limbic, and temporal cortices found in chimpanzees. In contrast, elderly rhesus monkeys show GM volume decline without the presence of neurofibrillary tangles (Alexander et al., 2008; Shamy et al., 2011). Taken together, regionally specific GM atrophy seems to be a common aspect of the primate brain aging pattern observed in macaque monkeys, chimpanzees, and humans. To make a case for the existence of Alzheimer’s disease in chimpanzees, validated cognitive tests for Alzheimer’s-like cognitive decline in non-human primates are needed, to test for direct associations between cognitive decline with tau pathology and brain atrophy.
To further analyze the possible moderator effects on aging, we considered the historical composition of the NCBR sample, with respect to the rearing environment. The majority of elderly chimpanzees over 40 years old (23/26) were born in the wild and captured at a young age, whereas only very few chimpanzees under 40 were wild born (5/168). The capture, separation from their mothers, and subsequent transport to the research centers can be considered a traumatic event with possible lasting effects on brain development and morphology (Bremner, 2006). In captivity, different chimpanzee-rearing experiences, either by their mother or in a nursery, has been shown to affect brain morphology (Bogart et al., 2014; Bard and Hopkins, 2018). The same should be expected in comparison of captive and wild-born chimpanzees. The disproportionate distribution of rearing and early life experiences likely influences our cross-sectional analyses of the effect of aging on GM volume. However, we have some reason to be confident that the aging effect shown here is not solely driven by these factors as rearing environment was added as a covariate to all age regression models and the VBM age regression model with and without rearing as a covariate are almost identical (Figure 5 and Figure 5—figure supplement 1 respectively). Moreover, the GM decline we found is extensive, widespread, and also present in chimpanzees under 30 years of age (p<0.0001), where 99% are captive born (143/144).
Hemispheric asymmetry was found in 68% (44/65) of all regions of the Davi130 parcellation, reproducing several regional findings reported in previous studies using diverse image processing methods as well as uncovering numerous novel population-level asymmetries. Previous studies utilizing a region-wise approach based on hand-drawn or atlas derived regions to analyze asymmetry in cortical thickness also reported leftward asymmetry of the insula (Hopkins et al., 2017) and rightward lateralization of cortical thickness of the PCC (Hopkins et al., 2017) as well as STG, MTG, and SMG (Hopkins and Avants, 2013). Previous VBM findings also revealed leftward asymmetry in the anterior SFG (Hopkins et al., 2008) along with rightward lateralization of the MFG, PrCG, PoCG, mOG, and CrusII (Hopkins et al., 2008; Hopkins and Avants, 2013). In the current study, new regions of larger GM volume in the left hemisphere were found in frontal (pMFG, mSFG, pSFG), temporal (aTTG), and parietal (PCun) cortices as well as in the basal ganglia (BF,GP, Pu). Novel rightward asymmetries could also be seen in the frontal (IFG, mOFC, FrOP), limbic (CC, Amy), temporal (pTTG, FFG), parietal (POP, AnG), and occipital (Calc) cortices besides the basal ganglia (Th, CN) and the cerebellum (CrusI, CerIX, CerVI, CerVIII).
The Davi130s’ region pTTG which contains the planum temporale (PT), presented significant rightward lateralization, while previous studies of the PT have shown leftward asymmetry in chimpanzee GM volume, surface area (Hopkins and Nir, 2010), and cytoarchitecture (Zilles et al., 1996; Gannon et al., 1998; Spocter et al., 2010). A possible reason for the divergence in this finding is that the anterior border of PT (Hopkins and Nir, 2010) lies several millimeters posterior from the anterior posterior split of the Davi130 TTG. Additionally, the left lateral sulcus in the Juna.Chimp template appears to proceed further posteriorly and superiorly compared to the right, which is consistent with previous findings in asymmetrical length of the POP in chimpanzees (Gilissen and Hopkins, 2013) and Sylvian fissure length in old world monkeys (Lyn et al., 2011; Marie et al., 2018). Population-level asymmetries in the pIFG in chimpanzees were documented almost two decades ago by Cantalupo and Hopkins, 2001, who reported a leftward asymmetry in pIFG volume in a small sample of great apes. In subsequent studies, this result could not be replicated when considering GM volume (Hopkins et al., 2008; Keller et al., 2009) or cytoarchitecture (Schenker et al., 2010). We also failed to find a leftward asymmetry in GM volume for the pIFG, in contrary to asymmetries found in humans (Amunts et al., 1999; Uylings et al., 2006; Keller et al., 2009).
A substantial amount of regions presenting significant inter-hemispheric differences of local morphology in chimpanzees has also been shown in humans (Good et al., 2001a; Plessen et al., 2014; Kong et al., 2018). Specifically, leftward lateralization has been found in human analogous regions of the SFG and insula utilizing GM thickness and volume in voxel-wise and atlas derived region-wise approaches (Good et al., 2001a; Takao et al., 2011; Plessen et al., 2014; Kong et al., 2018). Rightward asymmetry in the Davi130 regions IFG, STG, MTG, AnG, mOG, Calc, in addition to the thalamus and lateral cerebellum is documented in the human brain also using both VBM and surface measures (Good et al., 2001a; Takao et al., 2011; Plessen et al., 2014; Kong et al., 2018). Gross hemispheric asymmetry in humans follows a general structure of frontal rightward and occipital leftward asymmetry known as the ‘Yakovlevian torque’ (Toga and Thompson, 2003). This general organizational pattern of asymmetry was not apparent in the chimpanzee (Li et al., 2018).
The NCBR offers the largest and richest openly available dataset of chimpanzee brain MRI scans acquired over a decade with 1.5T and 3T MRI at two locations, capturing valuable inter-individual variation in one large heterogeneous sample. To account for the scanner effect on GM estimation, field strength was modeled as a covariate of no interest for analyzing the age effect on GM volume. The focus of this study was the analysis of GM volume, even though the CAT12 image processing pipeline enables surface projection and analysis. Consequently, the next step will be the application of CAT12 to analyze cortical surface area, curvature, gyrification, and thickness of the chimpanzee brain, to include behavioral data and the quantitative comparison to humans and other species, as cortical surface projection permits a direct inter-species comparison due to cross-species registration.
In conclusion, we present the new chimpanzee reference template Juna.Chimp, TPM’s, the Davi130 whole-brain parcellation, and the CAT12 preprocessing pipeline which is ready-to-use by the wider neuroimaging community. Investigations of age-related GM changes in chimpanzees using both region-wise and voxel-based morphometry showed substantial atrophy with age, which was also apparent in a matched human sample providing further evidence for human-like physiological aging processes in the chimpanzee brain. Examining population-based hemispheric asymmetry in chimpanzees showed a general rightward lateralization of higher GM volume without the presence of a distinct pattern like the ‘Yakovlevian torgue’ seen in humans.
This study analyzed structural T1w MRI scans of 223 chimpanzees (137 females; 9–54 y/o, mean age 26.9 ± 10.2 years, Figure 3—figure supplement 1) from the NCBR (http://www.chimpanzeebrain.org/). The chimpanzees were housed at two locations including, the National Center for Chimpanzee Care of The University of Texas MD Anderson Cancer Center (UTMDACC) and the Yerkes National Primate Research Center (YNPRC) of Emory University. The standard MR imaging procedures for chimpanzees at the YNPRC and UTMDACC are designed to minimize stress for the subjects. For an in-depth explanation of the imaging procedure please refer to Autrey et al., 2014. Seventy-six chimpanzees were scanned with a Siemens Trio 3 Tesla scanner (Siemens Medical Solutions USA, Inc, Malvern, Pennsylvania, USA). Most T1w images were collected using a three-dimensional gradient echo sequence with 0.6 × 0.6 × 0.6 resolution (pulse repetition = 2300 ms, echo time = 4.4 ms, number of signals averaged = 3). The remaining 147 chimpanzees were scanned using a 1.5T GE echo-speed Horizon LX MR scanner (GE Medical Systems, Milwaukee, WI), predominantly applying gradient echo sequence with 0.7 × 0.7 × 1.2 resolution (pulse repetition = 19.0 ms, echo time = 8.5 ms, number of signals averaged = 8).
The structural T1w images were provided by the NCBR in their original DICOM format and converted into Nifti using MRIcron (Rorden and Brett, 2000). If multiple scans were available, the average was computed. Following DICOM conversion, each image was cleaned of noise (Manjón et al., 2010) and signal inhomogeneity and resliced to 0.6 mm isotropic resolution. Finally, the anterior commissure was manually set as the center (0,0,0) of all Nifti’s to aid in affine preprocessing.
Structural image segmentation in CAT12 builds on the TPM-based approach employed by SPM12, whereby, the gray/white image intensity is aided with a priori tissue probabilities in initial segmentation and affine registration as it is in common template space. Another advantage of a TPM is that one has a template for initial affine registration, which then enables the segment maps to be non-linearly registered and spatially normalized to corresponding segment maps of the chimpanzee shooting templates. Lowering the possibility for registration errors improves the quality of the final normalized image. Improving upon SPM’s segmentation (Ashburner and Friston, 2005), CAT12 employs Local Adaptive Segmentation (LAS) (Dahnke et al., 2012), Adaptive Maximum A Posterior segmentation(AMAP) (Dahnke and Gaser, 2017; Gaser et al., 2020), and Partial Volume Estimation (PVE) (Tohka et al., 2004). LAS creates local intensity transformations for all tissue types to limit GM misclassification due to varying GM intensity in regions such as the occipital, basal ganglia, and motor cortex because of anatomical properties (e.g. high myelination and iron content). AMAP segmentation takes the initially segmented, aligned, and skull stripped image created utilizing the TPM and disregards the a priori information of the TPM, to conduct an adaptive AMAP estimation where local variations are modeled by slowly varying spatial functions (Rajapakse et al., 1997). Along with the classical three tissue types for segmentation (GM, WM, and CSF) based on the AMAP estimation, an additional two PVE classes (GM-WM and GM-CSF) are created resulting in an estimate of the fraction of each tissue type contained in each voxel. These features outlined above of our pipeline allow for more accurate tissue segmentation and therefore a better representation of macroanatomical GM levels for analysis.
An iterative process as by Franke et al., 2017 was employed to create the Juna.Chimp template, with T1 average, Shooting registration template (Ashburner and Friston, 2011), as well as the TPM (Figure 7). Initially, a first-generation template was produced using the ‘greater_ape’ template delivered by CAT (Franke et al., 2017; Gaser et al., 2020) that utilizes data provided in Rilling and Insel, 1999. The final segmentation takes the bias-corrected, intensity-normalized, and skull-stripped image together with the initial SPM-segmentation to conduct an AMAP estimation (Rajapakse et al., 1997) with a partial volume model for sub-voxel accuracy (Tohka et al., 2004). The affine normalized tissue segments of GM, white matter (WM), and cerebrospinal fluid (CSF) were used to create a new Shooting template that consists of four major non-linear normalization steps allowing to normalize new scans. To create a chimpanzee-specific TPM, we average the different Shooting template steps to benefit from the high spatial resolution of the final Shooting steps but also include the general affine aspects to avoid over-optimization. Besides the brain tissues the TPM also included two head tissues (bones and muscles) and a background class for standard SPM12 (Ashburner and Friston, 2005) and CAT12 preprocessing. An internal CAT atlas was written for each subject and mapped to the new chimpanzee template using the information from the Shooting registration. The CAT atlas maps were averaged by a median filter and finally manually corrected. This initial template was then used in the second iteration of CAT segmentation to establish the final chimpanzee-specific Juna.Chimp template, which was imported into the standard CAT12 preprocessing pipeline to create the final data used for the aging and asymmetry analyses.
The average T1 and final Shooting template were used for a manual delineation of macro-anatomical GM structures. Identification and annotation of major brain regions were performed manually using the program, 3D Slicer 4.10.1 (https://www.slicer.org). The labeling enables automated, region-based analysis of the entire chimpanzee brain and allows for robust statistical analysis. Nomenclature and location of regions were ascertained by consulting both chimpanzee and human brain atlases (Bailey and Bonin GV, 1950; Mai et al., 2015). The labeling was completed by two authors (S.V. and R.D.) and reviewed by two experts of chimpanzee brain anatomy (C.C.S. and W.D.H.). A total of 65 GM structures within the cerebrum and cerebellum of the left hemisphere were annotated and then flipped to the right hemisphere. The flipped annotations were then manually adapted to the morphology of the right hemisphere to have complete coverage of the chimpanzee brain with 130 labels.
The location of macroscopic brain regions was determined based on major gyri of the cerebral cortex, as well as distinct anatomical landmarks of the cerebellar cortex, and basal ganglia. Of note, the border between two adjacent gyri was set as the mid-point of the connecting sulcus, generally at the fundus. Large gyri were further subdivided into two or three parts based on their size and structural features to enable greater spatial resolution and better inter-regional comparison. Naming of regional subdivisions were based on spatial location, for example, anterior, middle, posterior, as these splits are based on macroanatomical features and do not necessarily correspond to functional parcellations.
Considering the limitations of macroscopic features present in T1w, we utilized distinct morphological representations to split large gyri, such as gyral/sulcal folds and continuation of sulci. If a distinguishable feature could not be determined, rough distance and regional size was employed as border defining criteria. The splits of the lateral temporal lobe, including the TTG, followed a continuation of the inferior portion of the postcentral sulcus that angles slightly posteriorly to better account for the increase in length of the gyri as it proceeds inferiorly. The central sulcus as well as the adjacent pre- and postcentral gyri contain a knob or U-shaped bend proceeding posteriorly. The superior beginning and inferior end of this bend were employed for the two splits of these gyri. Additionally, the central sulcus is the border between the frontal and parietal lobes, therefore, the FOP – POP split occurs at the termination of the central sulcus at the lateral fissure. Within the frontal cortex, the anterior posterior split of the MFG is at the meeting point of the middle frontal sulcus and the superior precentral sulcus, which translates to the inferior bend of the MFG. The tip of the fronto-orbito sulcus was used as an anchor point for the split of the pIFG and mIFG. The middle anterior split of the IFG was then determined by distance, whereby the remaining gyrus was separated into equally sized parts. The cingulate cortex anterior, middle, and posterior subdivisions were delineated by splits following the anterior and posterior bends of the gyrus around the corpus callosum. The cerebellum was divided into its major lobes which are quite similar across primates (Apps and Hawkes, 2009). Finally, splits within the OFC, SFG, and insula were based on equal size and/or distance.
CAT12 provides quality measures pertaining to noise, bias inhomogeneities, resolution and an overall compounded score of the original input image. Using these ratings, poor images were flagged for visual inspection when they were two standard deviations (std) away from the sample mean of each rating. The preprocessed modulated GM maps were then tested for sample inhomogeneity separately for each scanner (3T and 1.5T) and those that have a mean correlation below two std were flagged for visual inspection. Once the original image was flagged, affine GM, and modulated GM maps were inspected for poor quality, tissue misclassification, artefacts, irregular deformations, and very high intensities. For the second and third iteration, the passed modulated GM maps were tested again for mean correlation as a complete sample, flagging the images below two std for visual inspection, looking for the same features as in the initial QC iteration. Following the three iterations of QC a total of 194 of 223 chimpanzee MRI’s (130 females, 9–54 y/o, mean = 26.2 ± 9.9) qualified for statistical analysis.
A linear regression model was used to determine the effect of aging on total GM volume. Firstly, total GM volume for each subject was converted into a percentage of total intracranial volume (TIV) to account for the variation in head size. This was then entered into a linear regression model as the dependent variable with age, sex, scanner field strength, and rearing as the independents. Sex-specific models were conducted with males and females separately using age as the only dependent variable. The slope of each regression line was determined using R2 and a p-value of p≤0.05 was used to determine the significant effect of age and sex on total GM volume. The IXI brain development dataset (http://brain-development.org/ixi-dataset/) was utilized to compare the age effect on total GM volume between chimpanzees and humans, as it includes subjects with a wide age range and T1w images from MRI scanners of both 1.5T and 3T field strength. Prior to matching the IXI sample to the QC passed chimpanzee sample, all images collected from the Institute of Psychiatry (IOP) were removed to keep similarity to the chimpanzee sample of a single 1.5T scanner. After removing subjects without meta data, a total of 496 subjects (Figure 3—figure supplement 2) were used for matching to the chimpanzee sample regarding age, sex, and scanner field strength. To enable age matching between species, a factor of 1.5 of chimpanzee age was used to roughly calculate the comparable human age. This factor was chosen based on the comparable life span of the two species, because a chimpanzee 40+ years is considered elderly and so is a 60+ year old human, also a 60+ year old chimpanzee is very old and uncommon similarly to a human 90+ years old. Furthermore, the age of sexual maturity in humans is 19.5 years, while in chimps it is 13.5 years which is also approximately a difference of 1.5 (Robson and Wood, 2008). The sample matching was conducted using the ‘MatchIt’ (Ho et al., 2007) R package (https://cran.r-project.org/package=MatchIt) and utilizing the ‘optimal’ (Hansen and Klopfer, 2006) algorithm. The matched human sample contained 194 subjects (128 females, 20–78 y/o, mean = 39.4 ± 14.0) for statistical analysis.
The Davi130 parcellation was applied to the modulated GM maps to conduct region-wise morphometry analysis. First, the Davi130 regions were masked with a 0.1 GM mask to remove all non-GM portions of the regions. Subsequently, the average GM intensity of each region for all QC-passed chimpanzees was calculated. A multiple regression model was conducted for the labels from both hemispheres, whereby, the dependent variable was GM volume and the predictor variables were age, sex, TIV, scanner strength, and rearing. Significant age-related GM decline was established for a Davi130 label with a p≤0.05, after correcting for multiple comparisons using FWE (Holm, 1979).
VBM analysis was conducted using CAT12 to determine the effect of aging on local GM volume. The modulated and spatially normalized GM segments from each subject were spatially smoothed with a 4 mm FWHM (full width half maximum) kernel prior to analyses. To restrict the overall volume of interest, an implicit 0.4 GM mask was employed. As MRI field strength is known to influence image quality, and consequently, tissue classification, we included scanner strength in our VBM model as a covariate. The dependent variable in the model was age, with covariates of TIV, sex, scanner strength, and rearing. The VBM model was corrected for multiple comparisons using TFCE with 5000 permutations (Smith and Nichols, 2009). Significant clusters were determined at p≤0.05, after correcting for multiple comparisons using FWE.
As for the age regression analysis, all Davi130 parcels were masked with a 0.1 GM mask to remove non-GM portions within regions. Cortical hemispheric asymmetry of Davi130 labels was determined using the formula Asym = (L - R) / (L + R) * 0.5 (Kurth et al., 2015; Hopkins et al., 2017), whereby L and R represent the average GM volume for each region in the left and right hemisphere, respectively. Therefore, the bi-hemispheric Davi130 regions were converted into single Asym labels (n = 65) with positive Asym values indicating a leftward asymmetry, and negative values, a rightward bias. One-sample t-tests were conducted for each region under the null hypothesis of Asym = 0, and significant leftward or rightward asymmetry was determined with a p≤0.05, after correcting for multiple comparisons using FWE (Holm, 1979).
To illustrate the structural processing pipeline, we have created exemplar MATLAB SPM batch scripts that utilizes the Juna.Chimp templates in CAT12’s preprocessing workflow to conduct segmentation, spatial registration, and finally some basic age analysis on an openly available direct-to-download chimpanzee sample (http://www.chimpanzeebrain.org/). These scripts require the appropriate templates which can be downloaded from the Juna.Chimp web viewer (SPM/CAT_templates.zip) and then place the templates_animals/folder into the latest version CAT12 Toolbox directory (CAT12.7 r1609). The processing parameters are similar to those conducted in this study, although different DICOM conversions and denoising were conducted. Further information regarding each parameter can be viewed when opening the script in the SPM batch as well as the provided comments and README file. The code for the workflow in addition to the code used to conduct the aging effect and asymmetry analyses can be found here (https://github.com/viko18/JunaChimp; Vickery, 2020; copy archived at swh:1:rev:411f0610269416d4ee04eaf9670a9dc84e829ea0).
Age-related regional network of magnetic resonance imaging gray matter in the rhesus macaqueJournal of Neuroscience 28:2710–2718.https://doi.org/10.1523/JNEUROSCI.1852-07.2008
Imaging structural co-variance between human brain regionsNature Reviews Neuroscience 14:322–336.https://doi.org/10.1038/nrn3465
Broca's region revisited: cytoarchitecture and intersubject variabilityThe Journal of Comparative Neurology 412:319–341.https://doi.org/10.1002/(SICI)1096-9861(19990920)412:2<319::AID-CNE10>3.0.CO;2-7
Cerebellar cortical organization: a one-map hypothesisNature Reviews Neuroscience 10:670–681.https://doi.org/10.1038/nrn2698
The Isocortex of the ChimpanzeeUrbana: Univ of Illinois Press.
Local adaptive segmentationHum Brain Mapp Conference.
Good natured: the origins of right and wrong in humans and other animalsHarvard University Press.
Aged chimpanzees exhibit pathologic hallmarks of Alzheimer's diseaseNeurobiology of Aging 59:107–120.https://doi.org/10.1016/j.neurobiolaging.2017.07.006
Connectivity-based parcellation: critique and implicationsHuman Brain Mapping 36:4771–4792.https://doi.org/10.1002/hbm.22933
Premature brain aging in baboons resulting from moderate fetal undernutritionFrontiers in Aging Neuroscience 9:92.https://doi.org/10.3389/fnagi.2017.00092
Asymmetries in the Hippocampus and amygdala of chimpanzees (Pan Troglodytes)Behavioral Neuroscience 118:1460–1465.https://doi.org/10.1037/0735-7044.118.6.1460
Increased morphological asymmetry, evolvability and plasticity in human brain evolutionProceedings of the Royal Society B: Biological Sciences 280:20130575.https://doi.org/10.1098/rspb.2013.0575
Optimal full matching and related designs via network flowsJournal of Computational and Graphical Statistics 15:609–627.https://doi.org/10.1198/106186006X137047
A neuroanatomical predictor of mirror self-recognition in chimpanzeesSocial Cognitive and Affective Neuroscience 12:37–48.https://doi.org/10.1093/scan/nsw159
Brain weight throughout the life span of the chimpanzeeThe Journal of Comparative Neurology 409:567–572.https://doi.org/10.1002/(SICI)1096-9861(19990712)409:4<567::AID-CNE4>3.0.CO;2-J
Evolution of the central sulcus morphology in primatesBrain, Behavior and Evolution 84:19–30.https://doi.org/10.1159/000362431
Behavioral and brain asymmetries in primates: a preliminary evaluation of two evolutionary hypothesesAnnals of the New York Academy of Sciences 1359:65–83.https://doi.org/10.1111/nyas.12936
Regional and hemispheric variation in cortical thickness in chimpanzees (Pan Troglodytes)Journal of Neuroscience 33:5241–5248.https://doi.org/10.1523/JNEUROSCI.2996-12.2013
Imaging the evolution and pathophysiology of alzheimer diseaseNature Reviews Neuroscience 19:687–700.https://doi.org/10.1038/s41583-018-0067-3
Mapping brain asymmetry in health and disease through the ENIGMA consortiumHuman Brain Mapping 8:25033.https://doi.org/10.1002/hbm.25033
Atlas of the Human BrainElsevier.
Adaptive non-local means denoising of MR images with spatially varying noise levelsJournal of Magnetic Resonance Imaging 31:192–203.https://doi.org/10.1002/jmri.22003
A probabilistic atlas and reference system for the human brain: international consortium for brain mapping (ICBM)Philosophical Transactions of the Royal Society of London. Series B: Biological Sciences 356:1293–1322.https://doi.org/10.1098/rstb.2001.0915
Gray matter asymmetries in aging and neurodegeneration: a review and meta-analysisHuman Brain Mapping 38:5890–5904.https://doi.org/10.1002/hbm.23772
Statistical approach to segmentation of single-channel cerebral MR imagesIEEE Transactions on Medical Imaging 16:176–186.https://doi.org/10.1109/42.563663
The primate neocortex in comparative perspective using magnetic resonance imagingJournal of Human Evolution 37:191–223.https://doi.org/10.1006/jhev.1999.0313
Hominin life history: reconstruction and evolutionJournal of Anatomy 212:394–425.https://doi.org/10.1111/j.1469-7580.2008.00867.x
Ape Language: From Conditioned Response to SymbolOxford University Press.
Kanzi: The Ape at the Brink of the Human MindWiley.
Variability of broca’s area homologue in African great apes: Implications for language evolutionAnat Rec - Part a Discov Mol Cell Evol Biol 271:276–285.https://doi.org/10.1002/ar.a.10046
Animal Tool Behavior: The Use and Manufacture of Tools by AnimalsJohns Hopkins University Press.
Wernicke's area homologue in chimpanzees (Pan troglodytes) and its relation to the appearance of modern human languageProceedings of the Royal Society B: Biological Sciences 277:2165–2174.https://doi.org/10.1098/rspb.2010.0011
Primate CognitionOxford University Press.
JunaChimp, version swh:1:rev:411f0610269416d4ee04eaf9670a9dc84e829ea0Software Heritage.
Structural asymmetries in the human forebrain and the forebrain of Non-human primates and ratsNeuroscience & Biobehavioral Reviews 20:593–605.https://doi.org/10.1016/0149-7634(95)00072-0
Jonathan Erik PeelleReviewing Editor; Washington University in St. Louis, United States
Timothy E BehrensSenior Editor; University of Oxford, United Kingdom
Leah A KrubitzerReviewer; The University of California, Davis
Katherine L BryantReviewer; University of Oxford, United Kingdom
In the interests of transparency, eLife publishes the most substantive revision requests and the accompanying author responses.
This paper introduces a chimpanzee brain reference template using structural brain scans, along with the code and processing pipeline necessary for implementing the process. Using this framework the authors also observe age-related changes in gray matter and hemispheric asymmetry in chimpanzee brains that broadly mirror trends seen in human studies. The work represents an important step forward in comparative neuroanatomy and a useful resource for the field.
Decision letter after peer review:
Thank you for submitting your article "Chimpanzee Brain Morphometry Utilizing Standardized MRI Preprocessing and Macroanatomical Annotations" for consideration by eLife. Your article has been reviewed by three peer reviewers, and the evaluation has been overseen by a Reviewing Editor and Timothy Behrens as the Senior Editor. The following individuals involved in review of your submission have agreed to reveal their identity: Leah A Krubitzer (Reviewer #1); Katherine L Bryant (Reviewer #2).
The reviewers have discussed the reviews with one another and the Reviewing Editor has drafted this decision to help you prepare a revised submission.
We would like to draw your attention to changes in our revision policy that we have made in response to COVID-19 (https://elifesciences.org/articles/57162). Specifically, we are asking editors to accept without delay manuscripts, like yours, that they judge can stand as eLife papers without additional data, even if they feel that they would make the manuscript stronger. Thus the revisions requested below only address clarity and presentation.
Chimpanzees are our close relatives and their brains can help inform our understanding of human brain organization. Here a reference chimpanzee brain template is developed, and used to examine effects of aging and asymmetry. The atlas advances our understanding of the chimpanzee brain and is a valuable tool for comparative neuroanatomy.
Enthusiasm for the work was generally high. This is an important undertaking, and the transparent availability of the code was much appreciated.
1) If possible, it would be nice to see more detail regarding the comparative chimpanzee/human analyses (i.e., both symmetry and age-related decline). At minimum, some additional details in the text listing specific areas that are in common would be useful. If there is a way to link cross-species data in a quantified manner, that would be even better.
2) Understanding the limitations of the data, there was some concern about the use of sulcal boundaries and arbitrary subdivisions of the Davi130 parcellation. Are there other considerations that might be used in defining regions? Perhaps even explicitly addressing this issue would be useful. The parcellation is important not only for how we think about chimpanzee neuroanatomy, but as it directly effects the analyses and results. (This concern is helped somewhat by the voxel-based analyses.)https://doi.org/10.7554/eLife.60136.sa1
1) If possible, it would be nice to see more detail regarding the comparative chimpanzee/human analyses (i.e., both symmetry and age-related decline). At minimum, some additional details in the text listing specific areas that are in common would be useful. If there is a way to link cross-species data in a quantified manner, that would be even better.
We agree that more detail regarding the specific common regions that present age-related decline and GM lateralization will be beneficial for the reader to better understand the inter-species similarities and differences. Therefore, we have elaborated upon our human comparison in the Discussion and explicitly stated the comparable human – chimpanzee regions for each of the references we provide (see paragraph three and paragraph nine of the Discussion for aging and asymmetry respectively).
To address the point of a cross-species quantitative comparison we computed the age-related effect on global GM (Figure 3D) in a matched human sample. We choose the IXI MRI dataset (https://brain-development.org/ixi-dataset/) for comparison as it contains both 1.5T and 3T images with a large age range. The human sample was matched to the QC passed chimpanzee sample using the ‘MatchIt’ R package (https://cran.r-project.org/package=MatchIt) by matching age, sex, and scanner field strength optimal the ‘optimal’ matching technique. As life span and the aging process is different between species the human sample was match to 1.5* chimpanzee age (Robson et al., 2008). The matched human sample distribution is illustrated in Figure 3C. The analysis shows that the matched human sample presents a more prominent aging effect with less variance as compared to the chimpanzees. Ideally, we would register the chimpanzee GM volume aging effects to the human template space to directly compare the spatial distribution of age effects, but this inter-species volume transformation is by no means trivial. We are currently working on this problem but are not yet able to present a working implementation.
Aging, Discussion- “…Additionally, there was no significant difference in the age-related decline between humans and chimpanzee (Figure 3), even though a larger negative correlation with less variance was found in the matched human sample as compared to the chimpanzees demonstrates a commonality in the healthy aging process that was thought to be specific to humans. […] Similar age-related total GM decline along with presentation in homologous brain areas suggests common underlying neurophysiological processes in humans and chimpanzees due to shared primate evolution.”
Asymmetry, Discussion – “A substantial amount of regions presenting significant inter-hemispheric differences of local morphology in chimpanzees has also been shown in humans (Good et al., 2001a; Plessen et al., 2014; Kong et al., 2018). […] This general organizational pattern of asymmetry was not apparent in the chimpanzee (Li et al., 2018).”
2) Understanding the limitations of the data, there was some concern about the use of sulcal boundaries and arbitrary subdivisions of the Davi130 parcellation. Are there other considerations that might be used in defining regions? Perhaps even explicitly addressing this issue would be useful. The parcellation is important not only for how we think about chimpanzee neuroanatomy, but as it directly effects the analyses and results. (This concern is helped somewhat by the voxel-based analyses.)
The concerns about the boundaries and subdivisions of regions in the Davi130 parcellation are very much warranted as these delineations do seldomly align with known microanatomical boundaries (Amunts and Zilles, 2015). As we are constrained by the properties of the MRI acquisition method, especially the limited resolution of the T1-weighted images, we kept our regions relatively large. In general, we utilized gross macroanatomical landmarks when present for region delineation and subdivision such as, gyri and sulci peaks, fundi, and bends. We have now further elaborated on the exact landmarks we used for region subdivision in the Davi130 parcellation section of the Materials and methods. Moreover, we argue that the Davi130 parcellation, as a reduced representation of the chimpanzee brain based on macroanatomical features within a standardized space facilitates future inter-species and chimpanzee region-wise brain morphology and association to behavior (see paragraph two of the Discussion).
Subdivisions, Materials and methods – “Considering the limitations of macroscopic features present in T1w we utilized distinct morphological representations to split large gyri, such as gyral/sulcal folds and continuation of sulci. If a distinguishable feature could not be determined, rough distance and regional size was employed as border defining criteria. […] Finally, splits within the OFC, SFG, and insula were based on equal size and/or distance. “
Davi130, Discussion – ”Additionally, we provide the manually segmented, macro-anatomical Davi130 whole-brain parcellation comprising 130 cortical, sub-cortical and cerebellar brain regions, which enables systematic extraction of volumes-of-interest from chimpanzee MRI data. […] Furthermore, our macroscopic parcellation mitigates the curse of dimensionality for multivariate machine learning methods to be applied to the relatively small samples like the NCBR.”https://doi.org/10.7554/eLife.60136.sa2
- Sam Vickery
- Simon B Eickhoff
- Felix Hoffstaedter
- Sam Vickery
- Simon B Eickhoff
- Felix Hoffstaedter
- Svenja Caspers
- Svenja Caspers
- William D Hopkins
- Chet C Sherwood
- Chet C Sherwood
- Chet C Sherwood
- Steven J Schapiro
- Robert Dahnke
- William D Hopkins
- William D Hopkins
The funders had no role in study design, data collection and interpretation, or the decision to submit the work for publication.
We thank Jona Fischer for the creation of the interactive Juna.Chimp web viewer adapted from nehuba (github).
Animal experimentation: the chimpanzee imaging data were acquired under protocols approved by the Yerkes National Primate Research Center (YNPRC) at Emory University Institutional Animal Care and Use Committee (Approval number YER2001206).
- Timothy E Behrens, University of Oxford, United Kingdom
- Jonathan Erik Peelle, Washington University in St. Louis, United States
- Leah A Krubitzer, The University of California, Davis
- Katherine L Bryant, University of Oxford, United Kingdom
© 2020, Vickery et al.
This article is distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use and redistribution provided that the original author and source are credited.

domain: biology


The Mississippi Flexible Tax Incentive (MFlex), an economic development incentive created by economic developers across the state, will soon become law.
The Mississippi Senate took final action on Senate Bill 2159, authored by Senate Economic and Workforce Development Chairman David Parker, last week. The legislation now heads to the Governor’s Office for consideration.
“This program, which streamlines incentives to spur economic growth, has been several years in the making. The hard work of Chairman Parker and so many of our local economic developers have seen it through to fruition,” Lt. Governor Delbert Hosemann said. “As a performance-based—not promise-based—program, MFlex will make Mississippi even more competitive in the region.”
MFlex, which allows a credit against any state tax liability, combines several current incentives aimed at spurring economic development into one simplified option. The amount of the credit is calculated by considering the business’s investment in equipment and infrastructure, the number of full-time jobs created, and wages and benefits paid out to employees.
To participate in MFlex, a business must create at least 10 full-time jobs and make a capital investment of $2.5 million. Businesses qualifying for MFlex must report publicly the amount of investment, jobs created, average wage of employees, benefits provided, and other information.


text:
A leading Chinese fresh food delivery app has teamed up with Shanghai Ocean University to develop new seafood traceability standards.
Shanghai-based Ding Dong Maicai has launched its own seafood certification program, under which it will itself certify seafood sold on its platform on issues like water quality, feed input, and antibiotic residue contamination. The company made the announcement of the creation of Ding Dong Seafood G.A.P. in a newly-released green paper outlining its efforts to track its seafood products from farms to consumers’ doors.
It is not clear if the new standard being considered by Ding Dong is related to the Global G.A.P., a sustainability standard covering all aquaculture production systems for finfish, crustaceans and molluscs, covering legal compliance, food safety, workers’ occupational health and safety, animal welfare, and environmental and ecological care. SeafoodSource did not receive a response to emails sent requesting more information from Global G.A.P. or Ding Dong Maicai.
Ding Dong Maicai, which sells up to 10 tons of carp and perch and 10 tons of shrimp per day through its e-store and chain of offline stores, sources from 99 aquaculture suppliers in the Yangtze River delta, encompassing Shanghai and other prosperous cities like Hangzhou. Through the new program, its customers will now be able to access information on farm sources and feed inputs, Ding DongMaicai Head of Quality Control Lao Ai Jun said at the company’s official launch event for the program. Luo said the program, backed by assistance from researchers at Shanghai Ocean University, will specifically target the elimination of traces of chemicals like malachite green, chloramphenicol, and cadmium in the seafood sold through the company’s sales channels in order to give consumers confidence in food safety.
Ding Dong Maicai has raised USD 1 billion (EUR 820 million) to help it expand its network into northern China. It has experienced higher sales as Chinese consumers have turned to online food purchasing in greater numbers as a result of the COVID-19 pandemic. It announced on 9 June it is aiming for an initial public offering of stock in the U.S. this year in hopes of raising an additional USD 500 million (EUR 410.7 million), according to Reuters. Investors in Ding Dong include Sequoia Capital China and Tiger Global Management.
Ding Dong Maicai’s main competitors are Tencent-backed MissFresh, which also filed for a U.S. IPO on Tuesday, 8 June, and Meituan, which has raised USD 10 billion (EUR 8.2 billion) for expansion in the past year.
Photo courtesy of Andy Feng/Shutterstock

domain: food


text:
Bedford Tigers U13s have extended their unbeaten record this season after victory in Twickenham on Saturday.
Those from Bedford that had made the trip around the M25 saw the Tigers produce their best rugby league so far this season against Richmond, winning 46-14.
Built on the solid defence that the boys have shown all year they quite simply blew Richmond away scoring eight tries, converting seven and conceding just three.
The boys were dominant all over the pitch with strong running, solid tackling and great handling leaving Richmond with little to play for other than pride.
Try scorers were Luca Ventrone with two and Jack Harding and Matteen Querishi with three each. Toby Wiggett with the kicking duties was successful from the boot with seven out of eight.
One particular highlight was a prolonged period of defence on the Tigers line where Richmond tried to use their ‘battering rams’ to get over the try line from only five metres out only to be smashed back on each occasion by superb Tigers defence in particular led by Gurkeran Johal and Tom Pattinson.
This lasted until the pressure applied by the Tigers defence led to an error by Richmond and turnover ball.
This game saw the end of the current schedule of matches for the newly formed Bedford Tigers U13s.
Coached by Matt Wilson the squad has enjoyed great success winning all seven games.
The next match is scheduled as a curtain raiser at the Bedford Tigers Family Day on Saturday, August 6 at Wentworth Drive.
>> Bedford Tigers’ senior sides were both left without games on Saturday when North Herts pulled out of the fixtures.

domain: sports


text:
Choose your room
In the name of our entire staff, we welcome you happily to the Hotel Fahrenheit.
There are plenty of parking spots available to guests of this hotel.
Free for you
We provide guests with a complimentary bottle of water.
One of our hotel's courtesies is a free newspaper on a daily basis.
Make yourself at home
If you have any valuable items for safekeeping, please contact our qualified staff at reception or use the safe in your cosy room.
Our lift is available to take you to the upper floors of the cushy hotel.
We have video surveillance in our lobby.
Culinary greetings from the kitchen
Our vegetarian guests enjoy visiting our restaurant since it offers meals suitable to them. On the menu wholesome dishes are provided as well.
Where you will stay
Ask our experienced hotel staff about the easy-to-use air conditioning system that is part of each and every room.
At our hotel you'll also have a desk in your room.
There is also satellite TV provided in all of our rooms.
Our hotel also provides international newspapers in our rooms.
In all of the hotel rooms a convenient landline is available.
The bathrooms in this hotel have windows for your convenience.
Your hotel room's bathroom entails a shower. After you finish showering, grab a hot towel fresh from the heated towel holders and then dry your hair off with your hotel room's blow-dryer.
The rooms are equipped with sound proof windows.
The hotel offers many conveniences to make your stay at our hotel more enjoyable, for instance the sun studio.
Business hours of reception:
Every day: from 00:00 to 00:00

domain: travel