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"We have to be leaving, boys," Taylor and Sam's Mom called from the hallway.
They dropped their cereal bowls in the kitchen sink and picked up their backpacks on
the way out of the back door, slamming it shut.
A jar of blueberry jam and a jar of nutty peanut butter had been forgotten in the
morning rush, they were still sitting on the breakfast table instead of being put away in
the refrigerator.
"Blueberry," Nutty said, "I'm going to jump off this table and onto the floor, do you
want to come with me?"
"My glass jar might break when I land on the floor," Blueberry said.
"You won't break because I'll show you how to jump, watch me."
Nutty stood at the edge of the breakfast table and took a deep breath. "Here I go."
He spun himself around and twirled to the wooden kitchen floor.
"Did you watch me?" he said as he glanced up to see Blueberry looking over the edge of
the table.
"I did, but your jar is made of plastic and it won't break like mine."
"You have to be brave otherwise you'll never have any fun."
"Okay, you've talked me into trying it."
She looked around the kitchen for something to help her down instead of jumping off
the table like Nutty did.
There was a kitchen chair at the end of the table with a purple sweater hanging on the
side of it.
Blueberry twirled herself onto the chair and leaned on the sweater to slide down to the
wooden floor, landing beside Nutty.
"Phew! I made it."
"You did well. Now, let's go down the hallway."
They glided along the wooden floor until they came to the family dog, Julius. He's an all
white Husky with lots of fur.
"Grrr. What are you doing in my house?"
Blueberry shook with fear. "Um, we-we came from the k-k-kitchen table."
Nutty stayed quiet, he didn't like the dog's growling.
He had seen this dog's tongue licking his cousin's peanut butter jar clean.
Pumpkin, a Siamese cat with cream fur, dark brown ears and a very long tale, strolled
along the hallway to where Blueberry and Nutty stood by Julius.
She walked around them, sniffing.
"You both smell good, what are you?"
Blueberry's lid was loose and when Pumpkin sniffed her, she felt her lid tip a little.
"Nutty," she whispered, "is your lid screwed on tight? Mine isn't."
"It feels tight, but we may have a problem with yours.
Pumpkin is on her way back to you for another sniff."
Pumpkin's nose knocked Blueberry's lid right off.
"Oh! No, no. She's licking me! Yuck, stop that. Nutty, help me."
"Excuse us," Nutty said, "we have to be moving along."
"No," Pumpkin said, "you're not leaving just yet."
She turned to Julius. "You have to try this."
"I like peanut butter." Julius walked around Nutty's jar.
"We need to slide away from the big guys," Nutty whispered to Blueberry
"Soon, please."
Pumpkin and Julius argued about sharing the jars in front of them.
"I said, just a try," Pumpkin said. "I'm not letting you have half."
"Okay," Nutty said, "go, Blueberry, back to the kitchen."
They slid along as fast as they could go to an open cupboard door.
Pumpkin sniffed the wooden floor for Blueberry while Julius sniffed for Nutty.
There was only a slight smell because the two had slid away so fast.
In the far corner of the cupboard, Blueberry and Nutty clung to each other, praying the
cat and dog wouldn't find them.
"Pumpkin," Julius called, "they're around here somewhere. I can smell peanut butter."
"That means blueberry is close by as well." Pumpkin licked her lips. "Yum-yum."
Julius went to the breakfast table and stood with his paws on the edge of the table.
He spotted a knife with something on it that smelled really good. He dropped to the
floor, pulling the red and white chequered cloth with him.
Julius spotted the knife laying close by. "The knife," he said and then he carried it to the
corner and licked all the peanut butter off.
My, that was good.
"Where did you go?" Pumpkin asked. "I can't find Blueberry."
She turned to smell Julius and found peanut butter on his breath.
"You stinker, you found what you wanted but you didn't give me a single thought."
"That's not true. I found a little peanut butter on a knife, that's all."
"I'm getting bored looking for them. Good-bye."
"Okay, Pumpkin," he called after her as she jumped onto the counter to go out of the
open window. "I'll let you know if I find those tasty jars."
He turned back to the cupboard and pried the door open with his long nose.
"You two can come out of the cupboard now."
Blueberry and Nutty looked at each other in the semi-darkness.
Nutty whispered, "Do you believe him?"
"I'm too afraid to go out there until our family comes home."
"It's me Julius wants," Nutty said, "so I think I'm going to show my face and give him a
chase. I'll be back before you know it."
"Nutty, please be careful."
As Nutty stepped out of the cupboard, a car door slammed shut and kid's excited voices
could be heard at the back door.
]ulius glanced at the tempting peanut butter jar that was so close to him but decided the
family would be angry with him if they saw him chewing on it.
He pushed the jar with his nose to the tablecloth on the floor and left Nutty looking as if
he had fallen off the table when the cloth fell.
Taylor and Sam came rushing into the house feeling hungry. Sam opened the
refrigerator door looking for the jar of blueberry jam, but couldn't find it.
"Mom, where's the blueberry jam?"
"It was on the table this morning. It might still be there," She called from the next room.
Blueberry knew it was time for her to leave the safety of the cupboard.
She slipped though the open cupboard door and peaked around the corner of the
cabinets to see Nutty on the floor by Taylor's feet.
"What happened here?" Taylor asked, staring at the mess on the floor.
Blueberry slipped over to Nutty's side.
"Taylor," Sam called out. "The jar of jam it's by your feet, but it's missing its lid."
"What are you two doing on the floor and why is the table cloth on the floor?"
Taylor asked.
"Taylor," Mom said. "Are you talking to yourself?"
"Well, look at this mess. Who could have done this?"
"Your favourite dog, he's done it before.
Look what I found in the hallway, the blueberry jam's lid.
Last time, someone left the peanut butter jar's lid off he licked it clean."
"Julius wouldn't do anything like that."
"Remember, you only have to smell his breath for proof."
Julius stayed away from the kitchen while listening to Taylor telling Mom the story of
him being naughty. That is yummy memory.
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Hi MBC fam and welcome to our second week of the Artist Of The Week: Q&A series.
Chrisy Hurn who is the vocalist and guitarist of Basement Revolver reached out to me recently, to have a listen to her band's music and the possibility to publish their music on my blog.
Basement Revolver consists of the three members Chrisy Hurn (vocalist/guitarist), Brandon Monroe (Drums) and Nimal Agalawatte (Bass/Synth) who are from Hamilton, Ontario, Canada. Their genre is a mixture of Alternative/Indie Rock, Post-Rock, and Shoegaze.
Their music takes me back to the 90s with bands such as Garbage, Super Jesus, and Killing Heidi just to name a few.
So glad she reached out because this band is so talented!!!
And if you don't believe me, check out their latest single below.
Basement Revolver – Johnny Pt.2 (Audio)
Johnny Pt.2 is from the upcoming EP "Agatha". It is a continuation from their previous EP "Johnny". Agatha is due for release 21st July 2017.
To find out what the song is about, well you just have to read this exclusive interview to find out ;)
Chrisy talks about the band, past experiences, life, the meaning behind their latest song and album and what's next in store for the band in the coming months etc.
Please welcome my special guest Chrisy!!
Please tell the readers who are and where you're from?
Chrisy: We are from Hamilton, Ontario, Canada.
How old were each of you, when you knew you wanted to become an artist?
Chrisy: I was probably around 13 years old when I learned the guitar because I wanted to write songs.
Who was your inspiration, to get into music?
Chrisy: I am honestly not sure. I just started writing songs when I was thirteen, not really sure what brought it on.
What has been the biggest highlight throughout your career to date?
Chrisy: I think it has just been the ability to open for some bands that have meant a lot to us. One big one was when we opened for "We are the City". Overall, just the fact that we get to keep making music is a highlight.
From your experience in the music industry, what kind of advice would you give to other artists who want to get into the industry?
From your experience in the music industry, what kind of advice would you give to other artists who want to get into the industry?
Chrisy: To be honest we are just figuring it out for ourselves. Just keep working hard, and be kind - there is nothing worse than arrogance.
Your latest EP "Agatha", what was the process? And which song is your favourite?
Chrisy: The process is different for every song. Sometimes, songs come out of jamming with the full band (those are usually the best songs) and sometimes they come out of going through my old poetry notebooks. It is hard to pick a favourite because every song on the EP has been important to me for different reasons.
Johnny Pt.2 is the new single off the EP, what is the meaning behind it?
Chrisy: Johnny Pt. 2 is a continuation of Johnny. I wrote it when I was trying to move on from our breakup. There was still a lot I wanted to say and writing was the best way that I knew how to.
Your EP is to be released next month, what should the new and current fans expect?
Chrisy: Expect something a little different from last time, but not too different. All I can say is that we feel more confident in our sound than we used to and we are really happy with the outcome.
In the coming months, what should we look forward to?
Chrisy: A lot of things are in the works right now, most of them are secrets at this point, but expect to hear from us :)
And, what would you like to say to your fans?
Chrisy: Thank you so so much for your support. We wouldn't be anywhere without you, thank you for listening, sharing, and being excited about our music. We honestly are so thankful.
Thank you so much, Chrisy, for taking the time out of your busy schedule and for being a part of this awesome series.
To find out more about the band, make sure to follow, like and subscribe for the latest new music, news, tours and more by heading to the links below.
What makes this band stand out so much is their uniqueness, talent, light and shade, tone, expression and variety that they bring as a band and in their music.
Congrats on such a well-deserved career in the music industry, you definitely have what it takes to go further in this industry. I wish you all the best and look forward to seeing what's next in store now and in the future.
Make sure to purchase their new EP "Agatha" now on ITunes and you can listen to the album on Spotify via the links below.
*Pre-order new EP "Agatha" now on ITunes.
Release date: 21st July 2017
Thank you all so much for stopping by :)
I hope you enjoyed the second installment of this series. Stay tuned for more, exclusively at Music Beats Central!!!
And as always, don't forget to follow, like and subscribe for the latest new music, classics, interviews, reviews, news and more by heading to the links above or via the links below!
See you all soon in my next entry.
Have an amazing night/day everyone and talk soon!!!
Much love to you all and ciao for now!

subset: commoncrawl


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When I was in college, I took a class called the Biology of Aging. It was a wonderful course, with a considerate professor who was constantly reassuring and informing us about what was "normal" for our aging bodies. When it came to studying the integumentary system (skin, hair, nails and various glands located in the skin) Dr. B said, "The worst thing about skin is that it's so visible and people are judgmental." Ha! Ya think? But the wise doctor left us with deeper thoughts. He told us that in Eastern Europe they were proud of wrinkles. They tried to look older because that placed them into a higher pecking order. I'll bet that's why we often see pictures of an iconic Eastern European woman with a wrinkled suntanned face and a big huge smile. I believe I saw this type of picture during the Winter Olympics hosted by Russia. Obviously, people think differently in other parts of the world.
Suellen Ocean is co-author of The Common Sense Guide to Good Sex Available here:
eBook or computer download through Smashwords:
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eBook through Barnes & Noble:


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Eleven songs peaked on 2 February 1980.
Softly we met with a kiss
"Remember (Walking in the Sand)" by Aerosmith, a blues-rocking cover of the debut single by the Shangri-Las peaked at #67. Whereas the original sounds funereal, Aerosmith gave it a sharper bite by wisely dropping the moronic, naively maudlin vocal lead-in and making the chorus something of a rockabilly shuffle. It's still overdone, but less ridiculous, which succeeds in doing what a cover should do: reinterpret a song but not to the extent that it's no longer memorable. Not my cup of tea, but it serves its purpose.
Share my popcorn and jellybeans
After Prince's genre-defying "I Wanna Be Your Lover" we needed someone to remind us what mainstream disco sounds like, and Sister Sledge does so suitably with "Got to Love Somebody" (#64). Though it's straight up disco with the twonky bass, standard brass section, and far more singers than are necessary, Sister Sledge do better than the average disco group in the topics they sing about and the lyrics they use in doing so. This an empowering song that takes the specifics of loneliness without overselling loneliness as the end of the world– being the only hand in the popcorn box at the movies — and then the change in attitude that, presumably, will fix the situation. The song isn't making any promises other than that this girl is going to have fun looking for her next beau at the discos than she was watching rom-coms alone.
You've probably been crying forever
Sometimes you have to be careful with YouTube. I Don't Want to Talk About It" by Rod Stewart (#46) is a case in point: he rerecorded it in 1989, and I nearly reviewed the wrong version. This one is acoustic, and as a result feels a lot earthier, and more sincere, not adjectives I normally associate with Rod. His trademark gravelly voice works here to make him sound like he's on the verge of tears, a vulnerability I really appreciate in a good ballad. And it fits the lyrics: in consoling an ex who has been hurt in some way, he's absolving her, hinting strongly that he still loves her, sure, but not wanting to linger an the wrongs she's done him. The guitar work is nice, too (though I could have passed on the wonky key change toward the end), so all In all, a pleasant surprise.
Given any day there's a jet flying somewhere
John Stewart is a former member of The Kingston Trio, and given "Lost Her in the Sun" (#34), he must have been the one with the boring voice. Nevertheless, good songwriting and good delivery overcome vocal failings, and John Stewart delivers on this score. "Lost Her in the Sun" is an aching ballad about a lost love; he wonders what he's done that his girl should fly away without letting him know why, and he may never know. He does know it's going to hurt forever, like cold wind cutting deep into his soul. And he knows, whatever it was, it's his fault he's lost something wonderful — he's lost her in the warmth and light of the sun, after all. Really this song is about as perfect as a two-verse lost love song can get.
Dance with you, romance with you
Full disclosure: I was four years old in 1980, which means that like some of the young 'uns out there today who are unaware that Sting got his start in a band called The Police, I was unaware that Chaka Khan started out with Rufus (who isn't actually a person at all, but just the name of the band). They had a string of top-40 soul and disc hits through the 70s, of which "Do You Love What You Feel" (#30) was the final bookend. Lyrically, it's nothing special. Musically, it's fun, but not doing enough to really be memorable.
"Why Me" by Styx (#26) deserves its own entry.
Well, I wouldn't stop for a million bucks
"Don't Let Go" by Isaac Hayes (#18) is his last hit. It's a bit unusual for him in that it doesn't feature his voice the way you'd expect. He's pushed back in the mix, so much so that the jaunty funk guitar seems to get top billing over him. Don't get me wrong, this song is infectious; I dare you to listen to this without getting restless legs. It's just not a good showcase for Isaac Hayes. Really, this should be a Grace Jones song (a la "Pull Up to the Bumper", which fell one spot shy of the Hot 100 in 1981) — she can put the sultry sexiness that a choppy bouncy song needs, whereas Isaac Hayes is just too smooth for this kind of beat.
You're a different space in time
Speaking of Isaac Hayes, he's one of the talents behind "Deja Vu" by Dionne Warwick (#15); he wrote it with Adrienne Anderson, and Barry Manilow produced the whole album. Perhaps this is why the song sounds like living purple lame. Listening to it, can't you just imagine Dionne standing on a stage in, say, Las Vegas, wearing a purple lame gown, dripping in white rhinestones, exhaling this east ditty as a bunch of cigar-chomping businessmen sit around totally ignoring her? I say ignoring her, because the way she delivers this song is barely substantial; it's more a well-practiced breathing pattern than a series of words with natural inflection or even meaning. like so much cigar smoke and twinkling light, it drifts around in the background, greasing social skids but leaving not much of substance in its wake.
In the public eye, giving someone else a try
"Don't Do Me Like That" by Tom Petty & The Heartbreakers (#10) is another vulnerable rock song, albeit with a bit of a macho twang to it. Here's a guy who's trying hard not to admit he's in love with the girl he's seeing, and he's tryin' to play it all cool warning her if she strays she's going to get hurt, just as much as he would. It's a fine example of a guy transitioning from the free-wheeling womanizer to the marrying kind. I'm not sure exactly why this is top ten material, but the competition wasn't all that strong, as we've been seeing. If it sounds a bit on the high-school anthem side, like, oh, "Centerfold by the J. Geils Band, you're not alone: Wikipedia tells us that Tom Petty nearly gave this song to J. Geils, thinking it sounded more like their style than his.
All the debutantes in Houston
At #8, we have another loping bit of somnolent rocking from The Eagles, "The Long Run", which isn't making me like them any more than I did before. It's another rambling litany of related sentences that don't get much further than establishing that the singer was a cad, and now he's not, and it's because he's in love, and she should treat herself better, too. If it were a little more drunk it would sound just like "Heartache Tonight"; there may have been room on the charts for them both in 1980, but I certainly don't have the energy for both.
And finally, "Rock With You" by Michael Jackson spent its last week at #1, and every #1 deserves its own page.


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Monday, February 11, 2008
Job hunting... HELP!!
Well, it's THAT time again...
The debts are mounting and it's time to find another job... **grimaces**
It is, quite possibly, THE most scary thing right now as I'm aware I could end up absolutely anywhere and I've just got myself comfy in my current job. But the bank won't be too impressed if I carry on losing money, so it's off to the job centre I go...
My CV is sparkly new, I have an impressive new portfolio of stuff - now all I need is someone to give me a better paid job... So, if you know someone who is looking for a creative Copywriter with over seven years' experience, or you're a publisher looking for the next writing sensation, PLEASE get in touch!!
Well, it's worth a try, isn't it?! :o)
Posted by Miranda Dickinson at 2:05 PM
Labels: author, Copywriter, humour, jobs, publishers, scary, wombats, writer
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Tuesday, August 31, 2010
Sunday, August 29, 2010
End Of Summer Blues
Vintage Hat
Dress: Modcloths Chain Letter Dress
1950's Crinoline
Vintage Straw Bag: In my shop!
Shoes: Anthropologie, two years ago
I've been so excited about this post you guys! The dress I'm wearing is Modcloths Chain Letter Dress, an idea thought up by Starr of A Thought Is A Blossom. I'd been following the dress since Starr's post and all of the other fabulous ladies who got to style it, Rosie, Beatrice and Maria. So, when Maria of Lulu Letty asked me if I wanted to get the dress next I couldn't have been more excited! It was such a fun challenge to style the dress in my own way. So thanks Maria for making me a part of this fabulous idea! Who's the dress going to next? A lady who's pictures are always a dream and who calls Texas home, Leslie of Dalena Vintage!
I hope you all have had a lovely weekend! ;o)
Posted by Q's Daydream at 3:33 PM 33 comments
Friday, August 27, 2010
Pier Angeli, Like An Angel
I just recently discover Italian-born actress Anna Maria Pierangeli, or Pier Angeli. I was drawn to her timeless style and magical photos. I don't know much about Pier yet, I plan on ordering this book soon! I do know that we share the same birthday and James Dean was the love of her life. So in the meantime I'll share the gorgeous pictures with you and hope you find them as inspiring as I do.
Posted by Q's Daydream at 7:30 AM 21 comments
Tuesday, August 24, 2010
A Date With Fab
Soon after moving to Nashville I had the pleasure of meeting Fab of Vested Bee Vintage! Fab and I hit it off right away, and have become quick friends. The other day we got together with our hubbys in tow to take these pictures. We were quite the goofballs, so the serious pics really are tease...in between we were giggling and hopping about, hehe! This certainly won't be the last time you see Fab and I together! ;o)
Outfit Details
Blouse: New thrifted
Vintage Circle Skirt: In my shop!
Shoes: New thrifted
1960's Box Bag: In my shop!
Vintage 1930's Floral Dress
Vintage Bow Belt
Vintage Etienne Aigner Woven Bag
Shoes: UO
Posted by Q's Daydream at 6:00 PM 33 comments
Monday, August 23, 2010
Photo Shoot Sneak Peak
Posted by Q's Daydream at 6:43 AM 10 comments
Saturday, August 21, 2010
Sponsor Spotlight: Old Fashion Allies Vintage
I'd like to welcome me new sponsor Valerie of Old Fashion Allies Vintage! Old Fashion Allies is loaded with amazing vintage apparel, shoes and adorable collectibles! Here are just a few of my favorite things from Valerie's shop. Isn't that wedding dress fabulous?!
Posted by Q's Daydream at 1:36 PM 10 comments
Wednesday, August 18, 2010
A Dress Trade With An Elegant Gal
Vintage 1950's Dress: Trade with Casey
Belt: Made for me by Casey!
Purse: In my shop!
Shoes: Michael Kors, thrifted
A few weeks ago Casey and I decided it would be fun to do a vintage dress trade! Being that I adore Casey's style and we wear just about the same size I knew it would be the perfect match. When I received Casey's package in the mail I couldn't wait to tear that baby open, hehe! When I saw the dress I squealed with delight, then rushed to try it on. It couldn't have fit more perfectly! I'm madly in love with dress and I am going to wear it as much as I can while I have it, hehe. Not only did Casey send me this dream of a dress, but she made me the sweetest white bow belt that goes with it just wonderfully! She also surprised me with a great picture book of old Hollywood Star Shots and a mini pocket Elvis book! What's the next best thing to getting a new dress? It's doing a dress swap with a fabulously stylish gal! Thank you so much Casey! We should do this again sometime! ;o)
Be sure to check out Casey's blog for the dress I sent her!
Posted by Q's Daydream at 8:30 AM 39 comments
Sunday, August 15, 2010
Graceland, Beale Street & My Elvis Circle Skirt
Saturday morning at 5am Jonny and I got up and drove to Graceland! This past weekend was a big deal at Graceland. Every August they have what ya call Elvis Week, a celebration on Elvis's life. I have never been able to go to this and there was no way just being a few hours from Memphis I was going to miss it this year! While there we joined thousand of other Elvis fans of all ages in celebrating Elvis Presley. Me, I did it all in my Elvis circle skirt, hehe! So, here a just a few of the pics Jonny took...
All morning and afternoon we hung out at Graceland and took it all in. Here I am sitting at the Elvis fashion exhibit. It's a room filled with Elvis's personal wardrobe! *sigh*
Later that evening Jonny and I headed down to Beale Street and enjoyed some southern cooking and listening to the sounds of the blues, being played everywhere! I even got sung to by a sweet gentlemen on the street! After Beale Street we headed up the block just a bit to the gorgeous Orpheum theater to see a special screening of Elvis On Tour.
The movie was amazing! The audience was reacting as if Elvis was really on stage! Screaming, crying, clapping and laughing. Me I was quite, keeping my eyes on the screen the entire time. Later they showed never before seen rehearsal footage which was AMAZING! Then last but not least Elvis's background gospel group The Stamps came on stage and sang three songs. I think the entire audience had tears in they're eyes. Me, I sobbed! Hehe.
This is a screen capture Jonny snapped for me from a home movie. I think it's so sweet!
I hope you enjoyed my little recap of our Elvis adventures.
I'll talk to you all very soon! I have a fun photo shoot happening tomorrow with a new blogger friend and Wednesday a post I've been very excited about! :o)
Posted by Q's Daydream at 4:19 PM 14 comments
Friday, August 13, 2010
A Little Tribute...
Thirty-three years ago this Monday August 16th Elvis Presley passed away. You all know I'm a very big Elvis fan and I have been just about as long as I've been alive. Over the years I've seen every Elvis movie, heard almost every song and read just about every book. So, it almost feels like I knew Elvis. When I think of Elvis I honestly don't even think of him as being universally known and one of the biggest legends there is. To me he's just Elvis. Hopefully I don't sound to crazy to you all, haha! Anyway, I wanted to share with you the Elvis I love...
My favorite Elvis song...
Well my dears I'm off to Elvis Week! Pictures of my adventures when I get back. ;o)
Posted by Q's Daydream at 5:56 PM 10 comments
Etsy Update! Silk Sailor Dresses & Pretty Lace
Here are some of the things I've added to my shop today! My favorite has to be that amazing 1950's sailor dress. *sigh*
Posted by Q's Daydream at 3:05 PM 3 comments
Wednesday, August 11, 2010
Lena Hoschek+Dirndls=Fabulous!
These glorious photos of Lena Hoschek's 2010 Dirndl collection pretty much speak for themselves! Lena is constantly bringing us eye candy that you can't help but want to put on your body. I know I'm going to be in trouble when Lena's online shop opens it's doors! Happily in trouble, hehe! :o)
Posted by Q's Daydream at 6:50 AM 29 comments
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text:
Working with the Play! Framework means working with Akka, intentionally or not. But working with Akka Actors can be
tricky, especially when it comes to dependency injection. Play! 2.4 uses Google's
Guice for DI and of course it has the
ability to also bind Actors so an
ActorRef can be injected anywhere.
Single Actor DI
Biding and injecting one single Actor is simple and well documented . Just bind it in a Module:
package modules;
import actors.MyExampleActor;
import com.google.inject.AbstractModule;
import play.libs.akka.AkkaGuiceSupport;
public class ActorModule extends AbstractModule implements AkkaGuiceSupport {
public static final String EXAMPLE_ACTOR_DI_NAME = "my-example-actor";
protected void configure() {
bindActor(MyExampleActor.class, EXAMPLE_ACTOR_DI_NAME);
Enable the module in the app configuration:
play.modules.enabled = ${play.modules.enabled} [
Now you can inject the bound actor into every class using the
@Inject and
@Named annotation, e.g. in a Controller:
Keep in mind: Actors should always be injected as
ActorRef and not as the concrete class which has been
package controllers;
import akka.actor.ActorRef;
import akka.util.Timeout;
import modules.ActorModule;
import play.mvc.Controller;
import play.mvc.Result;
import scala.concurrent.Await;
import scala.concurrent.duration.Duration;
import scala.concurrent.duration.FiniteDuration;
import javax.inject.Inject;
import javax.inject.Named;
import java.util.concurrent.TimeUnit;
public class AssetController extends Controller {
ActorRef exampleActor;
public Result publish(String assetType, String assetUuid) {
FiniteDuration duration = Duration.create(5, TimeUnit.SECONDS);
String response = Await.result(
ask(exampleActor, "ask something", new Timeout(duration)).mapTo(classTag(Strijg.class)),
return ok(response);
// disclaimer: Stuff like error handling and message protocols intentionally left out in favor of clarity
Actor Pools
As you maybe know, there is the possibility to create pools of Actors instead of only one single instance. A pool behaves like a single actor, using a router to distribute messages to different child actors. The configuration options are huge, but most of the time, I personally only need an option to bind an actor pool using round robin distribution. No problems to achive this using the Akka configuration, but I could not find any documentation how to combine it using Play / Guice DI (in Java).
It turned out, that it is simple once you know it. Just pass the props of a
RoundRobinPool to the
bindActor method
as a third argument:
bindActor(MyExampleActor.class, EXAMPLE_ACTOR_DI_NAME, p -> new RoundRobinPool(5).props(p));
Now, instead of a single instance of
MyExampleActor there is a pool of 5 of them waiting for messages.
The example works for Play! Framework 2.4 using Java 8 and will maybe break in future versions.


text:
בטחון מוגבר בבי"ח רבקה זיו בעקבות תביעה בגין נסויים
בטחון מוגבר בבי"ח רבקה זיו בעקבות תביעה בגין נסויים
Uploaded by SilverRedIndigo on Mar 31, 2011
תושבי צפת ואזרחי מדינת ישראל יקרים
אם זה לא מדליק 1000 אורות אדומות הזהרה בעיניכם ומשמיע 1000 חצצרות התראה באוזנכם, אני לא יודעת מה יעשה את זה בשבילכם
The new security installations and other atypicals in Tzfat that should light a thousand red lights and blare a thousand trumpets that something terrible is happening in this town and it is being protected by the System.
The Hospital is now involved in a 30 Million Shekel joint lawsuit claiming human experimentation in contravention of the Geneva Convention.
I was able to get more shots of those those new "security" installations at the entrance of the Rivka Ziv Hospital in Tzfat (aka Safed and Zefat), Israel from the bus.
To remind you, just a few days ago; the story of a joint lawsuit against the Rivka Ziv Hospital to the tune of 30 Million Shekels broke in the news.
The Rivka Ziv Hospital admitted in a response to the Tzfat local paper חשיפה לצפון they they are indeed doing human experimentation, but insist they do it legally. At least one patient and the family of another patient who died claim otherwise.
The Management of the Hospital brazenly claims that they intend to expand their human experimentation as soon as the new Medical School about to be built in Tzfat is functioning.
Notice that the cameras are the same color as the iron bars over the Absorption Center for Ethiopian immigrants, which I got some footage of again in this video too.
Check out the "lights" at the Central Bus station in Tzfat.
And why are there two policemen in black uniforms "directing traffic" into the Rivka Ziv Hospital, when usually there are none.
Doreen Ellen Bell-Dotan, Tzfat, Israel


text:
The Mendicant Friars Necessary Today
Pope Benedict has dedicated these past Wednesday Audiences to teaching about the great mendicant friars, St. Francis and St. Dominic. These two pillars of renewal were called by the Spirit to transform the "dramatic and disquieting situation of the Church at that time, with her superficial faith that neither formed nor transformed life, her clergy little committed to its duties,…and the interior decay of her unity due to the rise of heretical movements".
At last week's Wednesday Audience Pope Benedict dedicated his catechesis to St. Francis of Assisi, referring to him as "a true giant of sanctity who continues to enthrall many people of all ages and religious beliefs". The Holy Father mentioned Pope Innocent III's dream in which he saw the Basilica of St. John Lateran about to collapse, but a 'small insignificant friar' held it up to prevent its fall. Pope Innocent recognized the friar to be Francis. "Innocent III was a powerful Pontiff, who possessed profound theological culture as well as great political power, but it was not he who renewed the Church. It was the 'small and insignificant' friar, it was Francis, called by God. Yet important to recall that Francis did not renew the Church without the Pope or against the Pope, but in communion with him. The two things went together: Peter's Successor, the bishops and the Church founded on apostolic succession, and the new charism that the Spirit had created at that moment to renew the Church".
At today's audience Pope Benedict heralded the life and legacy of Saint Dominic, founder of the Order of Preachers. Shortly before his birth in Caleruega, Spain in 1170, his mother, Bl. Juana of Osma, dreamed that a dog leapt from her womb carrying a torch in its mouth and seemed to set the world ablaze. Dominic "distinguished himself for his interest in the study of Sacred Scriptures and his love for the poor". After his ordiantion to the priesthood and election as a canon of the cathedral of Osma "he did not consider this as a personal privilege, nor as the first step in a brilliant ecclesiastical career; rather, as a service to be rendered with dedication and humility. Do not career and power represent a temptation to which even those who have roles of leadership and government in the Church are not immune?" the pope asked.
Upon traveling on diplomatic missions St. Dominic became aware of "the existence of peoples still un-evangelized,…and of the religious divides that weakened Christian life in the South of France, where activity of certain heretical groups created disturbance and distanced people from the truth of the faith". Dominic was asked by Pope Honorius III "to dedicate himself to preaching to the Albigensians" and he "enthusiastically accepted the mission, which he undertook through the example of his own life of poverty and austerity, through preaching the Gospel and through public discussions".
It was Dominic's itinerant apostolic life, rooted in "solid theological formation", "founded on Holy Scripture but respectful of the questions raised by reason" that sparked renewal in the Church. Pope Benedict teaches that "Christ is the most precious treasure that men and women of all times and places have the right to know and love! It is consoling to see how also in today's Church there are many people (pastors and lay faithful, members of ancient religious orders and of new ecclesial movements) who joyfully give their lives for the supreme ideal of announcing and bearing witness to the Gospel". He encouraged everyone, "pastors and lay people, to cultivate this 'cultural dimension' of the faith, that the beauty of Christian truth may be better understood and the faith truly nourished, strengthened and defended. In this Year for Priests, I invite seminarians and priests to respect the spiritual value of study. The quality of priestly ministry also depends on the generosity with which we apply ourselves to studying revealed truths".
Pope Benedict concluded, "With his sanctity, he shows us two indispensable means for making apostolic activity more incisive; firstly Marian Devotion, especially the praying of the Rosary which his spiritual children had the great merit of popularizing, and secondly, the value of prayers of intercession for the success of apostolic work".
Pray for all Franciscans and Dominicans, an increase in vocations to their consecrated life, and a renewed fidelity to their great founders' calling to make the most precious Treasure known, loved and defended.
Saint Francis and Saint Dominic pray for us!


text:
As 100 top chocolatiers from Africa, North and South America and Europe gather in Tübingen's idyllic old quarter to pay homage to the cocoa bean, chocolate lovers will be in their element. This is pure indulgence for all the senses, with melt-in-the-mouth confections, handmade gourmet chocolate, therapeutic chocolate massages, luxury chocolate tastings, creative praline courses, artistic cocoa paintings, inspiring readings and unusual chocolate theatre.
ChocolArt - Opening times:
03 Dec - 08 Dec 2019
Tue 1pm - 8pm
Wed - Fri 10am - 8pm
Sat 10am - 10pm
Sun 11am - 6pm
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text:
Ibiza 4 all
Ibiza History
Ibiza history
Ibiza History goes back to: 8 million years ago, at
the end of the Tertiary era, the Balearic islands, were
created as they arose from the ocean like all other European
mountain ranges.
In the beginning the islands were still attached to the Spanish mainland, but due to further rising of the sea level the Balearic Islands came about 12.000 years ago.
Ibiza was part of the Balearic Islands, and human habitation dates from at least 5000 BC.
Ibiza History tells us that the Balearic islands were positioned on the Phoenician trade route which soon caused the island being conquered and inhabited by Phoenicians, who named their new found port Ibossim, after their god Bes. They founded a settlement in Sa Caleta (the remnants are certainly worh a visit).
Back to the 9th century BC the Phoenician culture carried on trading lead, silver and minerals but due to covering great distances they needed places to lay in provisions. Ibiza very soon became their port of call where necessary supplies were put on board and repairs were carried out .
Later on the Carthaginians did the same but more important they also founded Ibiza Town, which became an important port in those days. They also used Ibiza as their burial grounds and constructed Les Salinas for saltmaking, Ibiza's most important trading product the 'White Gold' = Salt.
Archaeological discoveries have taught us much about the presence of the Carthaginians in Ibiza.
123 years BC, the island was occupied by the Romans, who, for the first time, governed Formentera, Ibiza (Ebusus), Majorca and Menorca as an archipelago. The islands most importantly supplied the Romans with necessary soldiers and wood while in the mean time they learned the local population how grow grapes, corn and olives.
Of course in those days. there were also constructed bridges, roads in the Balearic Islands, and cities such as Manacor, Palma, and the main town in those days: Pollentia now known as Alcudia.
Later on, in the 4th and 5th century, Ibiza was also occupied by the Vandals and Byzantines, and the first basilicas were established.
The Moors and the Vikings
came, but they were eventually displaced by the Arabs
who conquered the island in 707, and would remain for 5
centuries. Traces of all these sieges are clearly reflected
in the walled part of Ibiza -Upper Town, called D'Alt
Vila, meaning 'that high city'.
The ramparts of D Alt Vila include 7 watchtowers and 3 gates, including the Tablasgate - entrance gate, that are now declared unique historical monument in Europe.
Later in Ibiza History came, in
succession, the Romans, Visigoths and Muslims.
The Muslims left an important and recognizable imprint in
the island's typical architecture and the traditional
During Christian domination in the 13th century the Muslims were driven of the island by the Catalans, and then Ibiza quickly lost its importance for the region.
Once in 1479 Ferdinand of Aragon and Isabella of Castile got married, the Balearic Island officially belonged to the Kingdom of Spain and reconquest of the lost Spanish regions began.
This Bourbon government would stand until the 18th century.
The Balearic islands were now a major centre for maritime trade and thus a target for pirates from North Africa. Already for centuries Ibiza attracted the attention of pirates who were mainly specialized in stealing people. Formentera even was entirely depopulated by them.
Historians assume that the Ibicenco built their fincas
so far apart since previously larger settlements drew the attention of
the pirates at sea. The islanders thus painted their houses that pointed
towards the seafront in a more natural color that merged into the
The population of the island had been depleted by many wars and as a result the Ibicenco were extremely motivated to protect their island from further attacks from the sea. So they built protective watchtowers, each tower with views in different directions.
The news of approaching pirates was quickly passed through, from tower to tower, by signs of fire, so the population could find refuge in the protective churches that were already built.
The residents of Ibiza formed a sort of vigilante by acting
as pirates - corsair - themselves, in order to minimize the
attacks. Antonio Riquer Arabi in those days, was a
glorious corsair captain, which conquered more than 100
ships and helped prevent the island from much further
sufferings. To honor him the island later placed a memorial,
Corsair Obelisk, that still today is the only pirate statue
in the world.
In 1554, the now visible walls were built on top of the old city walls of arabic D'Alt Vila.
The town of D'Alt Vila, with watchtower at sea is also one true historic building dating from ancient times.
In the 17th century Punic dominaters chopped more than 4000 graves from the rocks, for their vaults.
The Archaeological Museum of Eivissa, at the Cathedral square in D Alt Vila, you can look at all survived Punic artefacts.
Remains of these times are found at:
- Es Soto: Punic water tanks, tombs belonging to the cemetery Puig des Molins, remains of an old Moslem cemetery and a Christian chapel
- Ses Feixes: historic irrigation system
- Las Salinas: historic salt flats & wetland of great beauty & ecological importance, with specific flora and fauna.
In the Museum of Ethnography of Eivissa in Can Ros des Puig the Missa in Santa Eulària des Riu, one can view collections of primitive objects and the Church of Jesús houses paintings from the historic Valencian School.
The imposing 13th century cathedral of Ibiza also houses an interesting museum.
Furthermore, Ibiza, also hosts a Museum of Modern Art for lovers of contemporary art.
In 1880 one begun construction of, what eventually would be,
180 windmills, that were used to pump water and for corn
grinding. Because of their importance for the rural
architecture of the Balearic Islands, although a large part
of the mills had lapsed, a number of them was now declared
to World Cultural Heritage by UNESCO such as Molí d'Toni
Joan, Molí des Porxet and Molí des Puig d'en
The nearly extinct Oceanic Posidonia - Seagrass fields in the water of the Oceanic Nature Reserve of Las Salinas was declared World heritage by UNESCO on december 14, 1999.
The mostly restored remains, from Ibiza's history of occupation, abandoned by various different prevailing cultures, are absolutely worthwhile visiting during a stay on this beautiful island, so you truly can imagine the life that the Ibizan population must have passed, during centuries of oppression by so many different cultures.
Even in the modern 20th century again there was excessive interest in Ibiza when in the 70-ties hippies came to the island and succeedingly clubbers in the 90-ties inevitably followed by the tourist industry.
An explanation can be found in the remarkable tolerance of the Ibicenzan inhabitants towards the annual invasion of exuberant and dissipated holidaymakers who only want to have fun no matter what.
As stated earlier, the local population is very tolerant to the sometimes odd-dressed holidaymakers, and strikingly extravagant gayscene that spread on their island in large numbers during the summer months.
The Ibicenco in this free, sunny island, live according to the motto, life and let live!
Don't leave Ibiza island without visiting the reminders of passed times of the Ibiza History.
- Via Romana, 31, 07800, Ibiza -
+34 971 301 771
1 April - 30 September
Tuesdays - Saturdays
10am - 2pm + 6pm - 8pm
Sunday 10am - 2pm
1 October - 31 March
Tuesdays - Saturdays 9am - 3pm
Sundays 10am - 2pm
Closed: Mondays & public holidays
Plaça de la Cathedral, 3 Dalt Vİla, 07800, Ibiza
+34 971 301 231
1 April - 30 September
Tuesdays - Saturdays
10am - 2pm + 6pm - 8pm
Sundays 10am - 2pm
1 October - 31 March
Tuesdays -Saturdays 9am - 3pm Sundays 10am - 2pm
Closed: Mondays & public holidays
Carrer Major, 2 Dalt Vila, 07800, Ibiza
+34 971 399 232
1 April -15 October,
Tuesdays - Saturdays
10am - 2pm + 5pm - 8pm,
Sundays 10am - 2pm
July & August,
Tuesdays - Saturdays
10am - 2pm + 6pm - 9pm
Sundays 10am - 2pm
16 October - 31 March,
Tuesdays - Saturdays 10am - 3pm Sundays 10am - 1pm
Ctra. Es Cubells, Cala Vadella, Cala d'Hort, Sant Josep de Sa Talaia, Ibiza
+34 971 338 154
1 April -30 September,
Tuesdays - Saturdays
10am - 2pm + 5.30pm - 9pm
1 October - 31 March
Tuesdays - Saturdays 10am - 2pm Closed: Sundays, Mondays, public holidays and from 20 December to 20 January.
Barrau Museum
Puig de Missa, 07840, Santa Eularia des Riu, Ibiza
+34 971 332 845
1 April -30 September,
Tuesdays - Saturdays
10am - 2pm + 5.30pm - 8pm
Sundays 11am - 1.30pm
1 October - 31 March
Mondays - Saturdays 10am - 2pm Sundays 11am - 1.30pm
Cathedral of Ibiza, Plaça de la Cathedral, 07800, Ibiza
Open: all year
Mondays - Saturdays 10am - 1pm month November
Wednesdays - Saturdays
10am - 1.30pm
Sundays Closed
Experience the feel of Ibiza's history, during the annual Eivissa Medieval Festival, (2nd weekend of May), held between the fortified walls of Dalt Vila. This festival commemorates the importance of the protective ancient fortification and its acknowledged World Heritage status by UNESCO in 1999. During the 4 day festival that attracts over 50.000 visitors, Dalt Vila, sets back in time by colorful decorations and breaths a Medieval atmosphere, where residents are dressed up in Medieval costumes, countless market stalls sell Medieval fare, craftsmen show their crafting skills, and endless entertainment & performances take place in a lively and unique Medieval festival atmosphere. (incl. free entry to all Dalt Villa's galleries & museums)
Various restaurants participate in EIVISSA GASTROMEDIEVAL, serving special Medieval Tapas & Menus on Thursdays, throughout May.
Ibiza History
Ibiza History - All about Ibiza History, most complete directory!


text:
Treat yourself like a royal guest in our Deluxe Suite. Located on the best spots of the hotel with the stunning view onto St Isaac's Cathedral will guarantee an unforgettable experience.
Our Deluxe Suite measures 50 sqm and comprises two beautifully appointed rooms. The bedroom features a king size bed. The elegant living room is an ideal place to relax after a day of viewing the St Petersburg's sights.
For more information about the Deluxe Suites, please feel free to contact us.


text:
Floral - Floral Print Shirt (Floral) | BARABAS
Leap into sunny days and warm weather in our bright FLORAL shirt. Demand attention with this comfortable and fashionable button down shirt.
- BARABAS Premium Shirt Collection
- Floral print
- Cotton/poly blend
- Collared
- Front button fastenings
- Long sleeved
Fabric & Care
- Fabric Content: 65% Cotton, 35% Polyester
- Care information: Machine wash at 40°C, do not tumble dry. Can be drycleaned.
- See sizing chart for exact measurements in Slim Fit and Classic Fit.


text:
of Contents
About the Author
Books by Bansi Pandit
What is Hindu Dharma?
Hindu View of
Why Hindus Worship
Principal Hindu
Law of Karma
Popular Systems
& Ethical
Ideals of Hindus
Hindu View ...
Hindu Reverence for
Daily Routine of a
Devout Hindu
Hindu Dharma
Hindu View of Ecology
Some Philosophical
Hindu Response
Contribution of
Practicing Hindu
Timeless Wisdom
Swãmi Vivekãnanda's
Works Cited
Color Plates
of Contents
The Hindu Temple Society of
North America
(Sri Maha Vallabha Ganapati Devasthanam)
Even though Swami Vivekãnanda
introduced and popularized the Hindu system of belief and philosophy in the
North American Continent in 1893, the movement of Hindu Temples began and has
gathered momentum only in the last quarter of this century. It is Lord Agastya,
the pre-eminent sage now living in the astral world, who took the initiative to
direct the establishment and growth of the New York Ganesha Temple and several
other major temples in this land.
Lord Agastya when invoked by sincere
devotees regularly through what is called Jîva Nãdî or Live Nãdî,
communicates with His devotees in the form of letters. These sacred letters
appear to the trained mediums only at the time of reading the sanctified ancient
palmyara leaf scriptures. In August 1968, Dr. Alagappa Alagappan, a
former U.N. Civil servant, was instructed to come and listen to the Nãdî
readings. On the first day, he was informed that Lord Ganesha would take an
abode in a city, the name of which began with the letter "N." When
asked to come on the following day, he was informed as to how this temple would
develop deep and abiding ties with the ancient heritage in India. The third time
he was summoned to come, he was instructed to take on the task of building the
Ganesha temple.
Every endeavor in Hinduism begins with
the worship of Ganesha and it was indeed appropriate that the first major temple
in this continent be built for Lord Ganesha. Dr. Alagappan proceeded to organize
a team under the leadership of Mr. C. V. Narasimhan and established the Hindu
Temple Society of North America on
January 26, 1970. Generous financial and technical support were provided by the
T.T. Devasthanam and the Endowments Department of the Government of Andhra
Pradesh, India. Literally thousands of devotees contributed their time and money
and the temple was consecrated by His Holiness Sri La Sri Pandrimalai Swamigal
July 4, 1977.
Since then the spectrum of worship has
broadened at this temple to include the major Hindu forms of worship and
practices prevailing in different parts of India. The Ganesha Temple which has
taken the name of Maha Vallabha Ganpati Devasthanam has drawn devotees in
ever widening concentric circles.
On September 21, 1995 more than eight
thousand devotees came unannounced and had the privilege of offering milk to
Lord Ganesha.
The New York Ganesha Temple has
actively participated in securing temple architects, artisans and priests for
several other temples in this continent. Since religion must be relevant to
daily life, every endeavor is made to organize activities which would contribute
to the welfare of devotees. Several cultural activities including instruction in
dance, music and Indian languages have been undertaken. The auditorium and Kalyana
Mandapam which will soon be in operation will provide some of the largest
facilities of their kind in New York. Over the years one of the main objectives
of the temple has been the organization of several growing social service
programs in India.
Recently a program has been launched to
disseminate substantive material pertaining to the Hindu religion and culture. A
one hour weekly TV broadcast was initiated on December 23, 1995. Another major
effort is to provide publications in simple and easily understandable English.
We are sponsoring publications such as
this for the benefit of a wider and young readership. Our biannual newsletter
which reaches more than 15 thousand readers also includes articles in popular
terms explaining the various aspects of Hinduism.
Dr. Uma Mysorekar
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text:
Riteish Deshmukh-starrer Banjo is likely to witness a good opening at the Indian as well as overseas box office on Friday, September 23. The movie has the potential to become Riteish's most successful solo release.
The actor has tasted success when it comes to multi-starrer films, the recent one being Housefull 3 that went on to enter the Rs 100 crore club. However, Riteish hasn't been fortunate enough with solo releases, except for the Marathi movie Lai Bhaari, which broke several regional box office records.
Meanwhile, the trailer and posters of Banjo have been creating a huge buzz. The widespread promotion, which lasted for more than two months, is expected to work in favour of the film. Despite getting released alongside two other films — Parched and Wah Taj — Banjo is expected to collect a decent amount at the Indian box office on its first day. However, it may face some challenge from last week's release, Pink, which has taken the box office by storm.
Riteish essays the role of a banjo player in the film and since music plays an integral part in the film, it is expected to draw music-driven audience to the theatres as well.
Set to be released in over 1,200 screens, Banjo is likely to get a boost from 5-10 percent advance booking on its opening day. The film, which also stars Nargis Fakri, is expected to collect over Rs 2 crore in the domestic market on its first day.
Banjo has a Maharashtrian favour to it and is hence expected to witness a bigger opening in the state as compared to other states. Also, although Riteish has a huge fan-base worldwide, his contribution to Marathi cinema and his performance in Lai Bhaari have made him a huge star in the state.
Banjo marks the Bollywood directorial debut of National-award winning Marathi director Ravi Jadhav. The film revolves around street artists, their issues and the challenges they face to create a niche for themselves in the world of music.


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Candida albicans undergoes a dramatic morphological transition in response to various growth conditions. This ability to switch from a yeast form to a hyphal form is required for its pathogenicity. The intractability of Candida to traditional genetic approaches has hampered the study of the molecular mechanism governing this developmental switch. Our approach is to use the more genetically tractable yeast Saccharomyces cerevisiae to yield clues about the molecular control of filamentation for further studies in Candida. G1 cyclins Cln1 and Cln2 have been implicated in the control of morphogenesis in S. cerevisiae. We show that C. albicans CLN1(CaCLN1) has the same cell cycle-specific expression pattern as CLN1 and CLN2 of S. cerevisiae. To investigate whether G1 cyclins are similarly involved in the regulation of cell morphogenesis during the yeast-to-hypha transition of C. albicans, we mutatedCaCLN1. Cacln1/Cacln1 cells were found to be slower than wild-type cells in cell cycle progression. TheCacln1/Cacln1 mutants were also defective in hyphal colony formation on several solid media. Furthermore, while mutant strains developed germ tubes under several hypha-inducing conditions, they were unable to maintain the hyphal growth mode in a synthetic hypha-inducing liquid medium and were deficient in the expression of hypha-specific genes in this medium. Our results suggest that CaCln1 may coordinately regulate hyphal development with signal transduction pathways in response to various environmental cues.
Candida albicans is the most prevalent fungal pathogen of humans. During infection,Candida undergoes a complex series of morphogenetic transitions, switching among a round, budding yeast form, a budding psuedohyphal form, and true tubular hyphae. The ability ofCandida cells to develop filamentous hyphae has been shown to contribute to its virulence: mutants that are defective in hyphal formation have been shown to have a greatly reduced level of killing in a mouse system, presumably because the hyphal form facilitates translocation across tissues and subsequent penetration (10, 17, 29, 37). Therefore, considerable effort has been directed toward understanding how the switch from yeast to hyphal cells inCandida is regulated. A number of environmental conditions are known to affect cell type switching in vitro; these include pH, temperature, and medium composition. Several genes specifically expressed in hyphal cells have been cloned, but relatively little is known about their regulation or the molecular mechanism governing the developmental switch (2, 6, 21, 53).
The recent observation that Saccharomyces cerevisiae also undergoes morphogenetic switching (20) has led to a new approach toward understanding filamentous growth in C. albicans. The experimental tractability of S. cerevisiae provides an important advantage over approaches used to study morphogenesis in Candida, a much less studied diploid asexual yeast. A number of specific genes and regulatory pathways have been shown to have a role in regulating filamentous growth inSaccharomyces. Based on this information, the homologs of some of these genes have been cloned and mutated in C. albicans and thereby shown to be involved in hyphal development (19, 24, 28, 34, 35, 37, 55). For example, elements of a conserved mitogen-activated protein (MAP) kinase pathway are required for pseudohyphal growth in S. cerevisiae (35, 41, 42, 44, 49). The corresponding components of the same MAP kinase pathway have also been shown to be involved in hyphal development inCandida (24, 28, 34, 35). Two transcriptional regulatory genes, C. albicans EFG1 (CaEFG1), whose homolog, the PHD1 gene, promotes pseudohyphal growth in S. cerevisiae, and the global transcriptional repressor gene C. albicans TUP1 (CaTUP1), have also recently been shown to be involved in regulating the morphogenetic switch in Candida (9, 55). Many other genes have been reported to affect pseudohyphal growth in Saccharomyces(3, 4, 7, 8, 18-20, 27, 36, 38, 39, 43, 48). By analogy to the pheromone-responsive MAP kinase pathway and to PHD1 andTUP1, many of these genes are likely to have homologs inCandida that function in hyphal development.
Studies done with S. cerevisiae have suggested that the transition between round yeast-form cells and long filamentous cells may involve regulation of the cyclin-dependent kinase (Cdk) system. InSaccharomyces, Cdc28 is the major Cdk that controls cell cycle progression at the G1/S-phase and the G2/M-phase transitions (45). Specific cyclin subunits bound to Cdc28 dictate the proper timing of cell cycle events, presumably by mediating its specificity. Manipulating the relative concentrations of G1 and G2 cyclins or the activity of the Cdc28 protein kinase affects the extent of polarized cell growth. Activation of Cdc28 by the G1 cyclins Cln1 and Cln2 (but not Cln3) promotes apical growth, while activation of Cdc28 by the mitotic cyclins Clb1 and Clb2 leads to isotropic growth (32, 33). Based on these and other results, Lew and Reed (32, 33) proposed a model in which cyclin-Cdk activities trigger different events in the morphogenesis cycle. Consistent with this model, the grr1 mutant, which stabilizes G1cyclins, also has enhanced filamentation (3, 4, 46). Furthermore, comparison of the cell cycle between the yeast form and the pseudohyphal form has revealed differences that implicate cyclins-Cdks in the regulation of filamentous growth. Unlike the yeast form, pseudohyphal cells divide symmetrically, have a shorter G1 phase and a longer G2 phase, and may possess an additional level of cell cycle control during G2(26). Taken together, these results suggest that the timing of the transition from G1 cyclin-Cdk predominance to G2 cyclin-Cdk predominance may be a regulated step involved in controlling the developmental switch between filamentous and yeast growth (25, 32).
In Saccharomyces, there are three major G1cyclins: Cln1, Cln2, and Cln3. While any of the three is sufficient to promote the onset of the cell cycle, they are not identical in function (13, 31, 56). Cln1 and Cln2 are more similar to each other, and their expression is strongly cell cycle regulated. Cln3 is only distantly related to Cln1 and Cln2, and its transcript level does not vary as dramatically through the cell cycle (45). Furthermore, the primary function of Cln3 is to activate the transcription of CLN1 and CLN2 through Swi4-Swi6 (56, 57), whereas Cln1 and Cln2 are much more potent activators of bud formation, DNA synthesis, and cell polarization (5, 12, 13, 33). To examine if G1 cyclins are involved in filamentous growth, we mutated the G1 cyclin genes in Saccharomyces. We found that diploid cln1 cln2 mutants are unable to form filamentous colonies on nitrogen starvation medium, while diploid cln3 mutants develop enhanced filaments (37a). Our genetic results suggest that Cln1 and Cln2 are involved in polarized cell growth during filamentation.
In order to determine whether G1 cyclins play a role inCandida hyphal development equivalent to that observed forSaccharomyces pseudohyphal growth, we determined the null phenotype of C. albicans CLN1 (CaCLN1). InC. albicans, two putative G1 cyclins have been isolated by their ability to complement a Saccharomycesstrain conditional for G1 cyclin activity or to confer resistance to pheromone-induced growth arrest upon overexpression (50, 58). However, the sequence similarities between these proteins and the Saccharomyces G1 cyclins are very low. In this report, we show that CaCLN1 mRNA has a periodic expression pattern similar to those of S. cerevisiae CLN1 and CLN2 mRNAs. We further address the role of Cln1 in filamentous growth by studying the phenotypes ofCacln1/Cacln1 null mutants. As predicted, theCacln1/Cacln1 mutants are somewhat slower than wild-type cells in cell cycle progression. We demonstrate that CaCln1 is necessary for the maintenance of hyphal growth on solid media and in liquid Lee's medium (30) but is not required for germ tube formation or hyphal growth in liquid serum-containing medium. Consistent with the morphological phenotypes, Cacln1/Cacln1strains have a medium-dependent impairment in the expression of hypha-specific genes.
Strains.The Candida strains used in this study are listed in Table 1. HLY strains were generated from CAI 4.
Plasmid isolation and construction. CaCLN1genomic DNA (4.5 kb) (pHL399) and CaCLN2 genomic DNA (4 kb) (pHL402) were isolated by colony hybridization of a Candidagenomic library in a Saccharomyces 2μm plasmid with PCR fragments of the respective coding sequences (34, 50, 58). The primers used for CaCLN1 had the sequences 5′CCGGAATTCCCATCCTCATACCATTCC and 5′CCGGAATTCCTGATTTATATTAACGTCAACGTC, and those used forCaCLN2 had the sequences 5′CCGGAATTCCTATCAATCCAAACATAGACAC and 5′CCGGAATTCGAAATGCAGAACATGATATTGTGG.
For CLN1 disruption (pHL419), a 1.3-kb upstream fragment that terminates at the SalI site at R23 and a 1.8-kb downstream fragment starting at the BclI site at L387 were cloned on either side of thehisG::URA3::hisGfusion in plasmid pMB7 (15). The linear fragment for integration was released by digestion with HindIII andKpnI before transformation.
For CLN1 complementation (pHL442), aBamHI/KpnI fragment from pHL399 was subcloned into a pBSII (Stratagene) vector in which the XbaI site had been destroyed. Then, the 5′ end of the insert was truncated to remove an XbaI site by releasing a BamHI/MluI fragment, followed by blunt-end ligation. Finally, a 1-kb PCR fragment containing the Candida URA3 gene was cloned into anXbaI site approximately 300 bp downstream of the termination codon of CLN1. The whole insert was then released byHpaI/KpnI digestion for transformation.
Microscopy. Candida cell morphology was photographed on a slide by use of a Zeiss Axioskop microscope with a ×100 or ×40 objective and Nomarski imaging. Candida colony morphology was photographed on solid media by use of a Zeiss Stemi 2000 microscope at about ×1.6 with dark-field imaging.
For staining with 4′,6-diamidino-2-phenylindole (DAPI) and Calcofluor,Candida cells were fixed in 70% ethanol and washed with 50 mM Tris–50 mM EDTA before use. For Candida cells in yeast form, DAPI and Calcofluor staining was performed as described previously (47). For Candida cells in hyphal form, prior to DAPI staining, cells were treated with Zymolyase (Sigma) at 25 μg/ml in phosphate-buffered saline for 30 min at 37°C to reduce the level of aggregation. Staining of hyphal cells with Calcofluor results in too much fluorescence all over the cell surface. To reduce the level of background staining, we treated cells with 50 μg of pronase and 20 μg of proteinase A per ml in phosphate-buffered saline for 2 h at 37°C before staining them with Calcofluor. Fluorescence microscopy was performed by use of a Zeiss Axioplan 2 microscope with a digital charge-coupled device camera.
Cell cultures. C. albicans strains were cultured essentially as described by Sherman et al. (51). Liquid Lee's medium (30) was modified by the substitution of 1% mannitol (Fisher) for glucose and by the dilution of all the other components by a factor of two (to improve solubility). Solid Lee's medium was prepared by the addition of Bacto Agar (Difco) to 1.5%. We prepared solid serum-containing medium by spreading 1 ml of newborn calf serum (Sigma) onto a 1.5% agar plate. Transformation ofCandida was performed by the method of Ito et al. (23), with the addition of 0.1 M dithiothreitol to the transformation mixture during polyethylene glycol incubation.
Cell synchronization.To isolate unbudded G1cells by centrifugal elutriation (14), Candidacells were grown to the early log phase in yeast extract-peptone–2% raffinose medium at 30°C. Raffinose was chosen as the carbon source because a higher percentage of cells are unbudded in a cycling culture grown in this medium than in YPD (glucose) medium (yeast extract-peptone-dextrose medium). Cells were collected by centrifugation, washed in ice-cold H2O, resuspended in cold H2O, sonicated to disperse clumps, and then loaded into the separation chamber of a Beckman JE-5.0 elutriation system maintained at 2,000 rpm. The pump pressure was gradually increased while the outflow was monitored microscopically. Unbudded cells were collected, concentrated by centrifugation, and then released into fresh prewarmed medium. Aliquots were removed periodically and centrifuged, and cell pellets were frozen in liquid N2.
Northern blotting.Total RNA was extracted from frozen cell pellets by phenol extraction (22). Formaldehyde gels were prepared and blotted essentially as described previously (22). DNA probes were labeled with a Stratagene Prime-It II random labeling kit and [α-32P]dCTP (DuPont NEN). An internal ClaI-HindIII fragment ofCaCLN1 and a ClaI-SalI fragment ofC. albicans ACT1 (CaACT1) were used as probes. Oligonucleotides with the sequences 5′CGGTCTTGACGTTGAAGATTCC and 5′CCTTTGGTACCATGACACCTGA were used to amplify a region outside the conserved cyclin box to probe for CaCLN2. PCRs were used to generate DNA fragments for probing C. albicans ECE1, C. albicans HWP1, and C. albicans HYR1(CaECE1, CaHWP1, and CaHYR1, respectively). The primers used had the sequences 5′TGACCGTGGAATTCAAGG and 5′GGACCATCTGCACCAGAAAGTG for HYR1, 5′GCCATCCACCATGCTCC and 5′GTGCTACTGAGCCGGCATCTC for ECE1, and 5′TGCTCCAGGTACTGAATCCGC and 5′GGCAGATGGTTGCATGAGTGG for HWP1. The sizes of mRNAs on our Northern blots correlated with the lengths expected based on information from theCandida Genome Database.
CaCLN1 is cell cycle regulated.To address whether regulation of the cell cycle is important for morphogenesis, we began to characterize G1 cyclin genes in Candida. We asked whether the previously reported Candida G1cyclin-like genes have the characteristic periodic cyclin mRNA expression pattern. In order to observe the periodicity ofCandida G1 cyclin expression, we purified synchronous populations of unbudded wild-type Candida cells by centrifugal elutriation and allowed the culture to reenter the cell cycle. Aliquots were removed from this synchronous culture at 10-min intervals, the percentage of budded cells was counted, and total RNA was prepared for RNA blotting (Fig. 1).
CaCLN1 and CaCLN2 genes had different patterns of expression during the cell cycle. CaCLN1 expression was periodic during the cell cycle. It peaked at 80 and 180 min after release, coinciding with the appearance of first and second buds. Bud emergence has been shown to occur as cells enter the S phase inCandida (52). Therefore, the time ofCaCLN1 expression corresponds to the G1/S-phase transition. At 140 min, expression was diminished, and at this point few cells had made a second bud; therefore, this time probably corresponds to the second G1 phase after release. Thus,CaCLN1 transcripts have a pattern of periodic expression during the cell cycle similar to that reported for S. cerevisiae CLN1 and CLN2. CaCLN2 showed some periodicity, but less than that of CaCLN1. In addition,CaCLN2 was detectable in very small unbudded cells.CaCLN2 accumulated rapidly after release, reaching a maximum at about 60 min. This time precedes bud formation and corresponds to the G1 phase. However, the transcript level was lower in the second cycle after release.
Disruption of the CaCLN1 gene.In order to determine whether G1 cyclins play a role inCandida hyphal development equivalent to that observed forSaccharomyces pseudohyphal growth, we investigated the null phenotype of G1 cyclin genes. Because CaCLN1 has a transcriptional pattern most similar to that of S. cerevisiae CLN1 and CLN2 and the S. cerevisiae cln1/cln1 cln2/cln2 mutant is the strain most defective in pseudohyphal growth (37a), we decided to focus on this gene. The genomic CaCLN1 gene was isolated, and a construct was made to interrupt the coding sequences by following the two-step knockout strategy of Fonzi et al. (15, 16) (Fig.2A). Both copies of CaCLN1were successfully disrupted in more than 10 isolates, which were derived from two independent heterozygous strains. Deletion of the internal portion of CaCLN1 was confirmed by PCR (Fig. 2B) and Southern blotting (data not shown). The loss of theCaCLN1 message in isolates with double deletions was also confirmed by Northern hybridization (data not shown). The growth rates of the heterozygous and homozygous Cacln1 mutant strains are somewhat lower than that of the wild type. At 37°C, the wild-type strain has a doubling time of 60 min in YPD medium, while theCacln1/Cacln1 mutant has a doubling time of 80 min. At 30°C, wild-type cells double every 70 min, whileCacln1/Cacln1 cells double every 90 min.
Cacln/Cacln1 cells are slightly retarded in cell cycle progression.To address whether there is any defect in cell cycle progression in the yeast form of the Cacln1/Cacln1 strain, we compared the timing of its cell cycle to that of the wild-type strain. We first determined daughter cell formation and nuclear division for Cacln1/Cacln1 and wild-type strains at 30°C. Synchronous cells were collected at 10-min intervals by releasing small unbudded cells prepared by elutriation into YPD medium at 30°C for yeast growth. Bud emergence and DAPI staining were used to measure daughter cell formation and nuclear division. The time from the maximally budded point of the first cell cycle after synchronization to the time of the maximally budded point of the second cell cycle is used as the time needed to complete one round of the cell cycle. In YPD medium at 30°C, wild-type cells took 80 min to complete one round, while mutant cells took about 100 min (Fig.3B, panels a and b). The slower cell cycle progression associated with the Cacln1/Cacln1 mutant was reproducibly observed in two independent elutriation experiments. Like Saccharomyces G1 cyclin cln1 cln2 and cln3 mutant cells (13), unbuddedCacln1/Cacln1 mutant cells purified by elutriation are slightly larger than wild-type cells. This larger cell size may explain why Cacln1/Cacln1 mutant cells entered the cell cycle earlier than wild-type cells, since cell size is known to be a critical determinant in cell cycle initiation in Saccharomyces(32).
The timing of the cell cycle for wild-type and mutant cells during the induction of the hyphal growth mode was examined. Small unbudded cells from the same elutriation experiment as that described above were released into YPD medium with 10% serum at 37°C. Both wild-type andCacln1/Cacln1 mutant cells were able to develop germ tubes under these conditions. Calcofluor was used to stain chitin ring structures (Fig. 3A). Since hyphal cells grow as a linear tube without any constrictions at each septum, bud emergence is difficult to define. Therefore, we used chitin ring formation as an indicator of hyphal septum formation. Chitin rings in yeast cells appear at the time of bud formation or immediately after bud emergence, indicating the G1/S-phase transition (32). DAPI staining was used to visualize nuclear division (Fig. 3A). Figure 3B, panels c and d, shows the percentage of hyphal septum formation and the percentage of cells that have completed nuclear division for synchronous cultures at 37°C in YPD medium with 10% serum. The time required for the completion of a cell cycle is calculated from the time at which 50% of cells have one chitin ring to the time at which 50% of cells have two chitin rings. The lengths of the cell cycle were about 80 min for wild-type cells and 110 min for mutant cells (Fig. 3B, panels c and d). Therefore, as at 30°C, mutant cells are slower in cell cycle progression at 37°C than wild-type cells. In this experiment, we observed that cells released into hyphal growth medium became asynchronous more rapidly than elutriated cells released into YPD medium. This finding may reflect an intrinsic feature of cell cycle regulation during hyphal growth.
The elutriation experiment with wild-type cells released into yeast and hyphal growth conditions showed that the length of the cell cycle for serum-induced hyphae was about the same as that for the yeast form. Chitin ring formation and nuclear division occurred at approximately the same time for yeast cells and hyphal cells. These data indicate that the cell cycle during germ tube formation is similar to that during yeast growth. However, this conclusion is necessarily limited to the specific hyphal induction condition used in this experiment. Furthermore, we can only examine the first two divisions after hyphal induction in this manner. Therefore, timing of the later cell cycles for the hyphal form may differ from that for the yeast form, as suggested by Kron and Gow (25).
Polarized apical growth precedes the periodic expression ofCaCLN1. Our Calcofluor labeling experiment shows that initial apical growth or germ tube formation preceded chitin ring formation (Fig. 3A). Small unbudded cells released into YPD medium at 30°C did not start to bud until 80 min, while cells released into YPD medium with 10% serum at 37°C initiated apical growth after 40 min. Chitin rings in hyphal cells formed at 80 min, about the same time as bud formation in yeast cells. Therefore, the formation of chitin rings lagged behind initial apical growth by 40 min during hyphal induction. The time of budding and hyphal septum formation coincided with the peak of CaCLN1 expression in several Northern hybridization experiments that we performed with synchronized cells released into serum-containing YPD medium at 37°C (Fig. 3C).
Hyphal growth is defective in Cacln1/Cacln1strains.The capacity of Cacln1 mutant strains to produce filaments was examined under many hypha-inducing conditions.Cacln1/Cacln1 strains have a profound defect in filamentous growth. The most dramatic effect is found on modified Lee's medium, a defined hypha-inducing medium containing moderate levels of nitrogen as ammonium, salts, amino acids, and mannitol as a carbon source (30). As shown in Fig. 4A, after 3 days on Lee's medium plates, the wild-type strain produces abundant filaments at the periphery of the colony, whileCacln1/Cacln1 and CaCLN1/Cacln1 mutant strains display none. After 5 days, some short filaments are observed around heterozygous colonies, but filaments are never observed around homozygous colonies. Furthermore, in contrast to the findings for wild-type cells, no filaments are observed under homozygous colonies after scraping or washing the mutant cells on the surface away (data not shown). We believe that the defect in hyphal development is probably not due to the lower growth rate of Cacln1/Cacln1strains because a longer incubation time did not allow the mutant strains to develop filaments. The phenomenon of a heterozygous mutant exhibiting a phenotype similar to that of the corresponding homozygote has been observed previously for Candida (24).
Cacln1 mutants also have a defect in the production of hyphal colonies on solid serum-containing medium (2% agar plated with 1 ml of serum). Both wild-type and Cacln1 mutant cells are capable of the initial response to serum, the production of long germ tubes (data not shown). However, after 1 day of incubation, the mutant strain begins to make predominantly round cells while the wild-type strain continues to grow exclusively as hyphae (data not shown). Figure4A shows growth on serum plates after 3 days of incubation. As it does on Lee's medium, the CaCLN1/Cacln1 strain exhibits an intermediate level of filament formation.
The ability of Cacln1 mutants to produce filaments was also tested with liquid media. We found little difference between mutant and wild-type strains in their ability to initiate germ tubes and produce elongated filaments in 5% newborn calf serum in YPD medium. Figure 4B shows hyphal cells after 4 h of serum induction at 37°C from overnight stationary-phase cultures. However, both wild-type andCacln1/Cacln1 mutant cells started to convert from hyphal growth form to yeast growth form after 6 to 8 h of incubation, and little difference in sustained hyphal formation was found between wild-type and mutant cells. However, we did observe that the mutant strain was slightly less clumpy during serum-induced hyphal formation than the wild-type strain (data not shown). In contrast to their behavior in liquid serum-containing medium, in liquid modified Lee's medium, Cacln1/Cacln1 strains were significantly impaired in the maintenance of the hyphal growth mode. Figure 4B shows the defect in filamentous growth after 15 h in Lee's medium. TheCacln1/Cacln1 mutant cells were able to initiate germ tubes and long cells at the beginning of the induction (data not shown), although the percentage of long cells was slightly lower in the mutant cells than in the wild-type cells. However, after prolonged incubation, the mutant cells mostly switched to a more pseudohyphal and then a yeast-form growth mode, whereas the wild-type cells continued to grow as more than 90% hyphal cells. Consistent with our other results, theCaCLN1/Cacln1 strain exhibited an intermediate phenotype.
To confirm that the defect in hyphal development observed in theCacln1/Cacln1 mutant was caused by CaCLN1disruption and not by other mutations introduced during two rounds of transformation, we replaced one of theCacln1::hisG alleles with a wild-typeCaCLN1 allele. Figure 5A is a schematic diagram of the Cacln1::CaCLN1complementation construct. Reintroduction of a wild-type copy ofCaCLN1 into the Cacln1 locus was confirmed by PCR and Southern blotting (data not shown). Two independentCacln1/Cacln1::CaCLN1 transformants were obtained, and both strains regained competence to produce hyphal colonies. The quality of the hyphae produced by the transformants was intermediate between that of wild-type and CaCLN1/Cacln1heterozygote strains on solid Lee's medium (Fig. 5B). The transformants also generated robust hyphal filaments in liquid Lee's medium, more similar to those of the wild-type strain than to those of the CaCLN1/Cacln1 heterozygote strain (data not shown). Therefore, reintroduction of a wild-type copy of CaCLN1 into the Cacln1/Cacln1 strain can complement the mutant defect in hyphal development.
Cacln1/Cacln1 mutants are defective in the hypha-specific transcriptional program.Because we observed a morphological defect for the Cacln1/Cacln1 strain in liquid Lee's medium, we began to wonder whether the expression of molecular markers for hyphal growth depends on CaCLN1. We examined the induction of three genes, HYR1, ECE1, andHWP1, known to be hypha specific (2, 6, 54). Overnight cultures were diluted 1:100 either into YPD medium for the yeast form of growth or into YPD medium plus 10% serum or Lee's medium for hyphal induction. Transcripts of these genes were undetectable in wild-type cells released into YPD medium at 30 or 37°C (Fig. 6). Furthermore, as expected, their mRNA levels increased dramatically within 60 min of the switch to the serum-containing medium. Transcriptional induction of hypha-specific genes in Lee's medium was slower. The level of transcripts at 3 h was generally lower than the 1-h induction seen in serum (Fig. 6). The levels of hypha-specific transcripts are consistent with the observed morphological effects. Germ tubes form immediately in response to serum, while hyphal development in Lee's medium starts after 3 h. The expression of hypha-specific genes in the Cacln1/Cacln1 mutant is consistent with the observed morphological phenotypes. The induction of HYR1,ECE1, and HWP1 in Cacln1/Cacln1 cells in response to serum is slightly reduced (about 50%) in comparison to that in wild-type cells (Fig. 6). In liquid Lee's medium, theCacln1/Cacln1 strain shows a profound defect in the transcriptional activation of all three hypha-specific genes. The difference in transcriptional induction between wild-type cells and mutant cells is even more dramatic after prolonged incubation. By 8 h of incubation in Lee's medium, wild-type cells have over several hundredfold the induction seen for CaECE1 andCaHWP1; the level of hypha-specific transcripts in the mutant cells remains at initial induction levels. The transcriptional defect in this strain precedes the detection of the morphological defect and is considerably more pronounced than the morphological impairment. This result suggests that G1 cyclins may play a relatively direct role in regulating the transcriptional program of hypha-specific genes.
G1 cyclins in Candida. S. cerevisiae has three major G1 cyclins. Cln1 and Cln2 are more homologous to each other in protein sequence, and their expression is cell cycle regulated, while Cln3 is only distantly related to Cln1 or Cln2, and its expression does not vary as dramatically through the cell cycle. Two putative G1cyclins, CaCln1 and CaCln2, have been cloned from C. albicans by functional complementation in Saccharomyces(50, 58). However, the protein sequence homologies between the Candida proteins and the SaccharomycesG1 cyclins are very low, with the highest similarity to Cln3 (50, 58). Our finding that CaCLN1transcripts are periodically expressed during the cell cycle suggests that CaCLN1 is a bona fide cyclin gene. Because its transcript levels peak at the time of bud formation, it is likely to be similar in function to the S. cerevisiae CLN1 andCLN2 genes. This notion agrees with the finding that the overexpression of CaCLN1 can cause pheromone resistance inSaccharomyces, as had been reported for S. cerevisiae CLN2 (11, 58). CaCLN2 shows a subtle periodicity in its transcription. CaCLN2 transcripts are expressed in unbudded G1 cells and peak earlier thanCaCLN1 transcripts. In addition, CaCLN2expression is lower in the second cell cycle. The pattern ofCaCLN2 expression is somewhat similar to that of S. cerevisiae CLN3 (57), suggesting that CaCLN2might be the counterpart of S. cerevisiae CLN3 inCandida.
The role of CaCln1 in filamentous growth. Cacln1/Cacln1mutants are unable to maintain the hyphal growth mode. This phenotype was observed on solid media and in liquid Lee's medium. The defect in hyphal colony formation agrees with our findings forSaccharomyces cln1 and cln2 mutants (37a). It also supports the previously reported finding that high levels of G1 cyclins promote polarized cell growth (3, 33). We believe that the defect in hyphal maintenance inCacln1/Cacln1 mutants is probably not due to its lower growth rate, because the mutant strains converted to yeast growth earlier than the wild-type strains. In addition, a longer incubation time did not allow the mutant strains to develop filaments.
Cacln1/Cacln1 mutants have immediately detectable defects in both cell cycle progression and transcription of hypha-specific genes. We found that Cacln1/Cacln1 mutant cells are slower in cell cycle progression than wild-type cells. Although we did not have a marker for the S phase in our experiments, it is possible that CaCln1 is involved in the timely progression of the cell cycle into the S phase in a manner analogous to that of S. cerevisiae Cln1 and Cln2. Our data also suggest that the G1 cyclin kinase may play a role in regulating the transcription of hypha-specific genes. We observed that Cacln1/Cacln1 mutants have rapidly arising defects in the transcription of several hypha-specific genes in both serum-containing medium and liquid Lee's medium and that this defect precedes the development of the morphological phenotype. The transcriptional defect is much more pronounced after longer incubation in Lee's medium. The defect in the transcriptional program of hypha-specific genes may account for part of the requirement for CaCln1 for the maintenance of hyphal growth. Several transcription factors, such as Tup1, Efg1, and Cph1, have been shown to regulate hyphal growth (9, 37, 55), suggesting that some of the hypha-specific genes regulated by these transcription factors are responsible for hyphal development. The three known hypha-specific genes,ECE1, HWP1, and HYR1, are not required for morphogenesis, as mutations in them do not affect filamentation (2, 6, 54). However, this fact may be due to the existence of other, functionally redundant genes, since genes with high sequence similarities to ECE1 and HYR1 have been discovered in the Candida Genome Sequencing Project. Another possibility is that other hypha-specific genes required for filamentation have not been discovered.
CaCln1 may have additional roles in filamentous growth, perhaps maintaining the polarization of the actin cytoskeleton, which is thought to contribute to the highly active apical growth at the hyphal tip (1). Like Saccharomyces cells,Candida cells in the yeast form of growth display a temporal change in the organization of the actin cytoskeleton during cell cycle progression (1). Studies with Saccharomyces have suggested that this temporal regulation of the actin cytoskeleton may be regulated by the Cdk Cdc28 (32). High G1cyclin-Cdk levels will result in a polarized actin cytoskeleton, and high B-type cyclin-Cdk levels will cause a depolarized actin cytoskeleton. In contrast to the situation during yeast growth, the majority of the actin cortical patches are concentrated at the tip of hyphal filaments during hyphal growth (1). Therefore, one role of CaCln1 in hyphal growth may be to maintain the polarization of the actin cytoskeleton. This activity of CaCln1 may be necessary for repressing the effect of the B-type cyclin-Cdk activity in the depolarization of the actin cytoskeleton in cycling cells (32) and is not required for polarized growth in cells with low B-type cyclin-Cdk activity. This notion is in agreement with our observation that stationary-phase Cacln1/Cacln1 cells are able to initiate apical growth when diluted into liquid Lee's medium. Similarly, initial apical growth from early G1 cells precedes hyphal septum formation (Fig. 3A and C) and the expression of cyclin genes. On the other hand, during prolonged hyphal growth post-Cdk activation, CaCln1 may be necessary to balance the effect of Cdk activities on the actin cyctoskeleton to prevent isotropic growth throughout the cell cycle.
Relationship between CaCln1 and signal transduction pathways in regulating hyphal development.We observed thatCacln1/Cacln1 cells are defective in hyphal development in liquid Lee's medium but that their hyphal growth is virtually unaffected in serum-containing liquid medium. This result suggests that two different signal transduction pathways may be responsible for hyphal development in serum-containing and Lee's media. Apparently, CaCln1 is more important for hyphal growth in Lee's medium than in serum-containing medium. This idea of two different signaling pathways is also in agreement with the fact that hyphal induction in serum-containing medium is much faster than that in Lee's medium. Several independent signal transduction pathways or regulatory proteins have been shown to be involved in hyphal development. These include a conserved MAP kinase pathway, an integrin-mediated pathway, and transcriptional factors, such as Tup1 and Efg1 (9, 17, 24, 28, 29, 34, 37, 55). Interestingly, the MAP kinase pathway and Efg1 have been mapped to two parallel pathways, because a double mutant blocks hyphal development under all growth conditions tested, while a single mutant does not (37). The CaCln1-Cdk kinase may function in parallel with these signal transduction pathways or as a potential target of one of the signal transduction pathways.
A crucial question concerning the role of cyclin-dependent kinase in filamentous growth is whether it is regulated in response to any extracellular signals. A careful comparison of budding and nuclear division between yeast growth and serum-induced hyphal growth did not reveal any changes in the timing of the cell cycle (Fig. 3B). However, our data cannot exclude the possibility that certain extracellular signals for hyphal development can induce the dimorphic switch by regulating the cell cycle machinery. Considering thatCandida cells can integrate a large number of extracellular signals and growth conditions as part of their dimorphic regulation, there are undoubtedly many mediators of these signals. Given the central role of the Cdk system in directing the localization of cell growth, it presents a likely target for some of these regulatory pathways.
We thank C. E. Birse for helpful discussions, S. B. Sandmeyer for comments on the manuscript, and anonymous reviewers for thoughtful critiques. We also thank W. Fonzi and J. F. Ernst for reagents.
This work was supported by NIH grant GM-55155 and by UC Universitywide AIDS Research Program grant K96-I-016 to H. Liu. H. Liu is a new investigator of Burroughs Wellcome Fund.
- Received 23 December 1998.
- Returned for modification 8 February 1999.
- Accepted 8 March 1999.
- Copyright © 1999 American Society for Microbiology


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