What are the most common bad habits of lacrosse goalies. How can goalies identify and correct these habits to improve their performance. Which techniques can help goalies overcome flinching and fear of the ball.
Understanding the Importance of Proper Technique for Lacrosse Goalies
Lacrosse goalies play a critical role in their team’s success, requiring a unique combination of physical skills, mental fortitude, and technical proficiency. The repetitive nature of their position makes it easy for bad habits to develop over time, potentially hindering their performance and growth as players. Recognizing and addressing these habits is crucial for goalies at all levels who want to elevate their game.
Why is proper technique so important for lacrosse goalies? Correct form and mechanics allow goalies to:
- React more quickly to shots
- Position themselves optimally in the goal
- Make saves more consistently
- Reduce the risk of injury
- Build confidence in their abilities
By identifying and correcting bad habits, goalies can unlock their full potential and become true assets to their teams. Let’s explore some of the most common pitfalls lacrosse goalies face and how to overcome them.
Overcoming the Fear of the Ball: A Critical First Step
Perhaps the most fundamental challenge many lacrosse goalies face is flinching or exhibiting fear when facing shots. This instinctive reaction can severely impact a goalie’s ability to make saves and must be addressed head-on.
How can goalies overcome their fear of the ball?
- Start with softer balls or tennis balls to build confidence
- Practice proper positioning to feel more protected
- Use visualization techniques to mentally prepare for shots
- Gradually increase shot speed and intensity in practice
- Focus on tracking the ball rather than anticipating impact
Remember, overcoming this fear takes time and consistent effort. Goalies should work closely with coaches and teammates to create a supportive environment for facing this challenge.
Correcting Stance and Positioning Errors
Proper stance and positioning form the foundation of effective goaltending in lacrosse. Many goalies develop habits that compromise their ability to cover the goal and react to shots efficiently.
Common stance and positioning mistakes include:
- Standing up too straight or hunching over excessively
- Keeping feet too close together or too far apart
- Failing to maintain a slight bend in the knees
- Holding the stick incorrectly or at an ineffective angle
- Positioning too deep in the goal or too far out
To correct these issues, goalies should focus on maintaining an athletic stance with feet shoulder-width apart, knees slightly bent, and body weight evenly distributed. Regular drills focusing on proper positioning and quick adjustments can help ingrain these correct habits.
Eliminating Premature Movement and Guessing
Anticipation is a valuable skill for lacrosse goalies, but it can become a liability when it leads to premature movement or guessing. These habits can leave goalies out of position and vulnerable to shots they might otherwise save.
How can goalies avoid the trap of premature movement?
- Train to react to the release of the shot rather than anticipating it
- Practice patience and discipline in maintaining proper positioning
- Develop better shot recognition skills to make informed decisions
- Work on explosive first-step movements to improve reaction time
By focusing on reading shooters and reacting appropriately, goalies can significantly improve their save percentage and overall effectiveness.
Improving Hand and Stick Placement for Better Saves
Proper hand and stick placement is crucial for making saves and controlling rebounds. Many goalies develop habits that leave them vulnerable to certain types of shots or limit their ability to direct the ball after a save.
Key aspects of hand and stick placement to focus on:
- Maintaining a relaxed grip on the stick
- Keeping hands in front of the body, ready to react
- Positioning the stick to cover the most dangerous shooting angles
- Practicing quick, precise movements to intercept shots
Goalies should incorporate drills specifically designed to improve hand-eye coordination and stick skills. Regular repetition of proper techniques will help ingrain these habits and make them second nature during games.
Addressing Communication and Leadership Shortcomings
Effective lacrosse goalies are more than just shot-stoppers; they are leaders on the field who direct their defense and contribute to overall team strategy. Many goalies underestimate the importance of this aspect of their role or struggle to communicate effectively during play.
How can goalies improve their communication and leadership skills?
- Develop a clear, loud voice for on-field instructions
- Learn and use standard defensive calls and terminology
- Practice directing defenders during drills and scrimmages
- Take responsibility for organizing the defense before each play
- Provide constructive feedback to teammates off the field
By embracing their role as a vocal leader, goalies can significantly enhance their value to the team and contribute to a stronger overall defensive unit.
Overcoming Mental Barriers and Building Confidence
The mental aspect of goaltending in lacrosse is often overlooked but can be just as crucial as physical skills. Many goalies struggle with confidence issues, negative self-talk, or difficulty bouncing back after giving up goals.
Strategies for improving mental toughness and confidence:
- Develop pre-game and in-game routines to stay focused
- Practice positive self-talk and visualization techniques
- Set realistic, achievable goals for each practice and game
- Learn to analyze mistakes constructively without dwelling on them
- Work with coaches or sports psychologists to address specific mental challenges
By strengthening their mental game, goalies can perform more consistently under pressure and bounce back more quickly from setbacks.
Enhancing Physical Conditioning and Athleticism
While often overlooked, a goalie’s physical condition plays a crucial role in their ability to perform at a high level. Many goalies neglect specific aspects of their physical training, leading to decreased performance and increased risk of injury.
What areas should lacrosse goalies focus on in their physical training?
- Improving overall cardiovascular fitness
- Developing explosive power for quick reactions
- Enhancing flexibility and mobility, especially in the hips and shoulders
- Strengthening core muscles for better balance and stability
- Incorporating plyometric exercises to improve reaction time
A well-rounded physical training program tailored to the specific demands of the goalie position can significantly improve performance and longevity in the sport.
By addressing these common bad habits and focusing on continuous improvement, lacrosse goalies can elevate their game to new heights. Remember, breaking bad habits takes time and consistent effort, but the results are well worth it. With dedication and proper guidance, goalies can transform their weaknesses into strengths and become true difference-makers for their teams.
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13 Bad Habits All Lacrosse Goalies Should Stop Immediately
The position of lacrosse goalie is one of repetition.
And for any sport that involves lots of repetition, it is very easy to develop bad habits.
Baseball pitchers who repeat the same throwing motion over and over suffer from bad habits on occasion just like lacrosse goalies who must repeat the same save movement over and over.
In working with youth goalies, the same bad habits tend to surface time and time again. Even more experienced goalies can develop some bad habits over time if they’re not careful.
The worst part is many lacrosse goalies don’t even realize they have these bad habits. That’s where the power of a good lacrosse coach comes into play. They can help spot and correct these bad habits.
But even being aware of bad lacrosse goalie habits can be half the battle in getting them corrected.
Flinching or being scared of the ball is by far the single worst and most common habit a goalie can have. However this action is not often voluntary.
I discussed remedies for flinching, false steps, and general fright of the ball in my post 4 Common Lacrosse Goalies Problems and How to Fix Them.
Today I’m going to write about bad lacrosse goalie habits that are completely voluntary and can be fixed with the right coaching, the right preparation, and the right drills.
As lacrosse goalies our goalie in making a save is to make a single, precise movement to the shot.
When we drop our hands on high shots, it’s a double movement. Hands drop and then do back up.
Now you still may make the save like the young keeper does in the GIF above but as the players get better and the shots get faster, this bad habit will ruin you.
This habit is so common in young goalies (older ones too for that matter) that I wrote an entire post on how to stop dropping your hands on the shot.
When I first warmup a keeper, without giving any instruction I’ll analyze their tendencies and see what needs to be fixed.
Some goalies upon making a save will throw the ball back to the coach and then turn to look at the pipe to get setup in their ready position.
This means looking away from the ball. Bad!
As a goalie we never want to look away from ball. We should be concentrating on the ball at all times.
Getting the goalie to break this habit of looking away from the ball to check his position will vastly improve their concentration.
There is enough going on on the lacrosse field. The goalie doesn’t need to distract himself by taking his/her eyes off of the ball.
This one is similar to the first item. Some goalies will make marks in the dirt or tape on the turf within the crease so they know the right positions on their arc.
But guess what? To use them, you have to look down!
And as we just learned looking away from the ball is a bad habit that needs to be broken.
If the goalie is struggling to find the right spot on their arc, you can do two things:
1. Landmark Technique
Teach your goalie to pick out large features of the landscape around the field such a tree or a building.
Select one for left center (45 degree) and right center (45 degree). For top center you can obviously use the other goal.
As the game is played, these landmarks can be viewed in the background without focus. This way the goalies doesn’t drop the eyes and take focus off the ball but still maintains good position in between the pipes.
2. Switch to a Flat Arc
One of the benefits of playing a flat arc is that its easier to get setup in the right position.
If your goalie struggles with maintaining good positioning when moving through the different spots on his arc perhaps consider trying a flatter arc.
Most goals are made by moving the goalie out of good position so setting up in the right place on the arc is half the battle when trying to make saves.
Besides saving shots the lax goalie has many other responsibilities.
They’re the leader of the defense and need to call out the right lacrosse defensive terms to ensure their team knows whats going on.
However, many times these other responsibilities get in the way of the goalie’s primary responsibility: stop the shot!
Goalies must be ready for shots as soon as the ball enters their defensive side.
If you have the right ready stance, it shouldn’t be uncomfortable to stay in it for a full 5 minute possession by the other team.
If it is tiring, that’s a clear sign that you need to hit the gym to build strength in your shoulder, glute, and leg muscles.
There’s really no excuse for this one and yet I see it happen all the time, even at the top levels. Goalies need to be ready for every shot!
Notice in the photo below the shooter is about ready to rip one and yet the goalie is not ready. He’s checking is pipe, from his feet it looks like he’s moving. Nothing compared to what I discuss in the perfect lacrosse goalies stance.
Analyze your goalie (or yourself) during practice or games when the shots are being released. Are you in a good ready position for 100% of the shots you face?
If you’re not ready for the shot, break that habit immediately.
Always be ready, even in practice drills.
There are actually some respected lacrosse goalie coaches (like Trevor Tierney shown above) that teach this technique where the keeper drops to their knees on low shots.
But I consider this a bad habit. I never coach my goalies to drop to their knees on low shots.
You will make more saves over time staying on your feet.
In the event of a rebound you’re better prepared to make a second save if you stay on your feet. Furthermore, going to your knees often produces more rebounds because the goalie cannot get his body over the ball to control the bounces.
Going down to your knees on the save limits your ability to make outlet passes. Many times one of your middies will have slipped behind the offense and by staying on our feet during a save can we make that quick outlet pass to start the fast break.
That outlet pass may only be there for a split second and in the time the keeper goes from knees back to feet the opportunity may be gone.
Also remember you only have 4 precious seconds to throw an outlet pass or leave the crease once you gain possession. Getting back up from your knees to your feet eats up one of those valuable seconds.
Finally its seems that for goalies who’ve been taught to drop their knees, this becomes their default action for every shot especially when they’re nervous. They’re dropping to their knees instead of reacting to the shot. This is horrible as smart attackman will simply start shooting high and score goal after goal on you.
Therefore if the goalie has developed the habit of dropping to his/her knees on low shots this needs to be broken.
Be sure to check out my post on tips for saving low shots.
If you’ve read this blog for awhile you should understand the basic mechanics of making a save.
Our lead foot takes a step to whatever side the ball is shot on. Many goalies have no problem with this step however they forget to follow with a trail step.
As a lacrosse goalie we want to get our entire body behind the ball during a save.
The trail step, which is taken with opposite foot that took the lead step, will help ensure our body is behind the ball during a save.
If you’re not taking a trail step you’re also not exploding off of your back foot to attack the ball.
The trail step allows us to finish the save in a nice and even balanced position, ready to make a quick outlet pass or another save if there was a rebound.
In this GIF of professional lacrosse goalie Kip Turner of the Chesapeake Bayhawks we see that he takes his lead step with his left foot. Then watch the trail step with his right foot to ensure his body stays behind the ball. He ends this save process in a very balanced position, the exact purpose of our trial step.
If you’re goalie is not finishing his/her saves with a trail step, you can do the drill called “The Glyde” that I describe in my post on lacrosse goalie drills.
This drill helps goalies focus on completing the save process with a trail step.
A lacrosse goalie’s body should always stay square to the shooter.
Pre-shot, during the save process, and post shot our body should remain square to the shooter.
Many goalies have a bad habit of twisting or rotating their bodies while making a save. This is especially prevalent for the off-stick saves where you have to rotate the stick to meet the ball.
The problem with rotating our torso is that:
1.) We reduce the surface area of body. Our chests have much more surface area than the side of the body. So by keeping our body square to the shooter we remain “big” in the goal. Which is to say we take up the most surface area possible.
2.) Risk injury. Goalies wear pads on their chest, not on the sides of body. Therefore, a goalie who has a bad habit of turning their body on saves is subjecting themselves to an unnecessary injury risk. Keep your body square to the shooter and a shot that happens to get by your stick will hit you in the chest pad, not in the exposed ribs.
While warming up the goalie or during drills point out when the goalie turns his body and attempt to correct it immediately.
You can also point out these moments while reviewing video with your goalie.
If you’re looking for a specific lacrosse goalie drill to help keepers maintain their torso square to the shooter on shots, try the Football Drill described in this post.
Never turn your body like this during a save. Always remain square to the shooter.
A common bad habit especially among newer goalies is not talking enough on defense.
The goalie is a leader of the defense and a leader who doesn’t communicate hurts the team.
I wrote a full post on how to get your defense and goalie to talk more.
The summary version is this.
1.) Make sure you know what to say
Many times lack of communication stems from the goalie’s lack of knowledge of the defense. Review all the defensive terms and start using them in practice.
2.) Use names
In the chaos that can be a lacrosse game your teammates are going to hear you better if you use their name.
People respond better to their names.
3.) Be Loud in Practice
If you’re a naturally shy or quiet person, you’re going to have to break that habit. You have to get outside your comfort zone. When I was a shy sophomore our coach used to put me in a circle with the entire team surrounding me and make me yell stupid things at the top of my lungs.
Whatever it takes to break out of your comfort zone and break the habit of not communicating to your defense.
The goalie needs to be the loudest player on the field.
Even if you know all the proper defensive terms to use, if your team can’t hear them, what’s the point?
A goalie’s communication needs to be loud, clear, and with the proper tone.
For a great example of how loud a goalie needs to be checkout former Loyola Greyhounds goalie Jack Runkel communicating and encouraging his defense in a game against Duke.
If your goalie has the bad habit of not communicating loud enough be sure to emphasize this throughout the practice.
When you’re standing on the sideline, can you hear the goalie’s commands loud and clear? If not, he/she needs to improve their voice volume.
You can also have them perform some voice strengthening drills to develop a stronger voice that can be heard by the entire team.
Making a save in the sport of lacrosse is hard. That’s why even top goalies only have a save percentage between 50-60%
So when a goalie does make a save and then makes a lazy outlet pass for a turnover it drives a coach absolutely nuts.
There’s no excuse for lazy outlet passes.
Throwing lazy outlet passes is mostly mental. The goalie after making the save relaxes and doesn’t put his/her full effort into a very critical aspect of goalie’s games: leading the clear.
So assuming that the goalie knows how to throw proper outlet passes, coach them to remain mentally alert after the save and put 100% effort into every pass.
Many goalies have the bad habit of what I call “happy feet”.
When we’re setup in our lacrosse stance we want to be absolutely still.
Happy feet is when a goalie is moving his feet or even body or hands while waiting for the shot to come.
I’m not talking about moving from one spot to another while on your goalie arc. That movement is required.
But once you’re in the right spot on your arc you should be still. If you’re moving it could cost you a 1/2 step or a fraction of a second to then move in the right direction of the ball.
And as we’ve learned, especially at the upper levels, a fraction of a second is the difference between a goal and a save.
When our body is still we’re ready to explode in any direction to save a shot.
Please don’t confuse being still with not being in an athletic stance. As we learned in the perfect lacrosse goalie stance, we’re always in an athletic position. This bad habit simply refers to excess and unnecessary movement.
This bad habit is somewhat inherent and a goalie might not even realize they’re doing it.
Videotaping a goalie during practice and showing him/her the bad habit can help them identify when they might be going into ‘happy feet’ mode.
Also repeating a key word like “STILL” might help the goalie focus on remaining still while setup in their ready stance.
The idea of lacrosse saves is that we get our body behind the ball. This isn’t like an ice hockey goalie where we attempt to make kick saves.
Therefore if a lacrosse goalie is continually kicking their leg out during saves to either side of their body this is a bad habit that must be changed.
Why? Because when we’re kicking out leg out we’re not stepping to get in front of the shot. We’re not driving off our back leg to get our body and stick in the right place to make a save.
I’ll admit it makes for a dramatic photo hence the reason we see it on the cover of Lacrosse Magazine. But notice in the cover pic how this goalie’s body is not behind that shot. If the ball misses her stick, or hits the top and ricochets, that’s a goal for sure.
If you notice yourself or your goalie kicking instead of driving off their back foot and stepping to the ball, be sure to correct this bad habit.
The final bad habit I see lacrosse goalies endulge in all the time has to do more on the mental side of this position.
It’s negative self-talk.
If you had a teammate who talked to you the way you talk to yourself, would you want to be around that person?
Pay attention to your self-talk. For most of us, it’s horrible. Things like:
- I’m a horrible goalie
- We lost that game because of me
- Our team would better if someone else was the goalie
Make an effort to stop any and all negative self-talk, and replace it with something positive.
Have your own back.
To be an All-American lacrosse goalie you must identify and eliminate all bad habits.
By first understanding these common bad habits for lax goalies we can ensure that we’re not doing these behaviors on the field.
As a coach you’ll want to look for these bad habits and correct them as soon as possible.
With practice, recognition, and repetition there’s no reason any lacrosse goalie should continue with these 11 bad habits because they’re 100% voluntary.
Until next time! Coach Damon
Any bad lacrosse goalie habits you have that I missed? Let me know in the comments.
Berkeley Girls Head Coach Steps Down – Tampa Lax Report
The Berkeley Buccaneers varsity girls lacrosse head coach stepped down from her position last week after adopting a son and wanting to spend more time with her new family.
Coach Jennifer Stoll and her husband Adam adopted the boy in December and Coach Stoll found that the extra time spent coaching, doing film work, and all the coaching duties were keeping her from fully enjoying their new son.
“We had been anxiously awaiting Ben’s arrival for several years and the moment we met him, our whole world changed” Stoll said.
“I was hopeful that I could balance my roles as a new mom, a Lower Division teacher at Berkeley, and the Head Varsity Coach of a lacrosse program I have had the honor of being a part of since 2010.”
Stoll continued, “Missing bedtimes with late night games and rushing home from practice to get a few minutes with Ben was challenging.”
In Coach Stoll’s message to her players, she lamented the thought of the program she built since 2010 suffering due to her new family duties. “I do not want the program to suffer due to changes in my personal life. I want the program to continue to grow and flourish. Because of this, I think it is time for someone else to have the opportunity to lead the program – someone who can put all they have into making the program even better.”
Berkeley Prep Athletic Director Bobby Reinhart was thankful and appreciative of the program Stoll has fostered. “Jen Stoll launched Berkeley girls lacrosse and put our program on the map” he said. “From simple and modest beginnings Jen has built a program that is now incredibly competitive.
Jen is a great coach who has earned the admiration and respect of every girl she has coached. Jen will be greatly missed in our athletic program. We wish Jen nothing but the best in this new and special chapter of her life.”
Stoll will continue to teach at Berkeley Prep as well as support the girls lacrosse program as she is able.
Berkeley is currently looking to fill this vacancy. Interested applicants should contact Athletic Director Bobby Reinhart via email – [email protected].
Denison’s Mike Caravana to step down as head men’s lacrosse coach
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GRANVILLE, Ohio— The 2021 Denison men’s lacrosse season came to an end on May 15 after another run into the NCAA postseason. On Sunday morning longtime head coach Mike Caravana announced to his team that he would be stepping away from the program and passing the torch to the next steward of Denison Lacrosse.
If you spent some time chatting with Coach C during his 28-year tenure leading Denison lacrosse, he was always quick to point out that he was merely a caretaker of a program that was far greater than himself. Taking the reins in 1991 from Tommy Thomsen, this ethos provided the backbone for one of the most historically successful programs in Division III sports.
Putting others above yourself. Playing for the person next to you. Having the personal pride and drive to improve not only your game but all aspects of your life. These were just a few of the lessons that Caravana worked to instill in hundreds of Denison lacrosse alumni through the years.
“It has been an honor to be a part of the Denison community for all of these years. I take great pride in all that we have been able to accomplish over the nearly three decades, both on and off the field,” noted Caravana. “I am so grateful for the dedication and commitment made by all my players and assistant coaches through the years.
Our continued high level of success could not have occurred without them and their hard work. Their support and generosity, enthusiasm, and loyalty reassure me that my years here were both impactful and fulfilling.”
Caravana arrived at Denison in the fall of 1990 and was hired to not only replace the fabled Thomsen but he was also asked to serve as the head coach of the Denison men’s soccer team, a position he held until 1996. Caravana continued at Denison until 2005 when a new opportunity at Woodberry Forest School called him and his family to the Virginia boarding school. Three years later, the Denison head coaching position was vacant and Caravana returned to the Hill.
Since he re-assumed the role as head coach in 2009, Denison has become a staple in the Division III national championship conversation, winning 85 percent of its games while advancing to the NCAA quarterfinals six times and the NCAA Semifinals in 2017.
Across all levels of NCAA lacrosse, Caravana ranks 18th in winning percentage and total wins.
He is just one of 21 coaches in college lacrosse history to reach the 300-win milestone. He ranks 10th in Division III history in wins (320) and seventh in winning percentage (767). In 1994 he was named the USILA Division III Coach of the Year and he is a seven-time North Coast Athletic Conference Coach of the Year with his most recent award coming in 2021.
Denison has won 15 NCAC Championships and six of the eight NCAC Tournaments since it was introduced in 2013. If you’re talking NCAA Tournaments you might as well just pencil the Big Red into your bracket at the start of every season. Eighteen times Caravana has taken his squad to the NCAA postseason with a remarkable 11 appearances in the quarterfinals and Final Four berths in 1999, 2001, and 2017.
The people underneath the helmets are Caravana’s greatest achievement. He’s coached 75 USILA All-Americans, 17 USILA Scholar All-Americans, three first-team All-Americans, one NCAA Postgraduate Scholar, 15 NCAC Players of the Year and 133 first-team All-Conference picks.
Caravana has never had a losing season at Denison and in 2016 he passed his predecessor, Thomsen, as Denison’s all-time leader in wins.
Professional development is also part of Caravana’s path. Not only did he enter as the head coach of two sports at Denison, from 2003-06 he served as an assistant coach with the United States Lacrosse National Team that resulted in a Silver medal at the International Lacrosse Federation World Championship. In 2016, Caravana also spent one season as the acting head field hockey coach while current head coach, PJ Soteriades, was out on leave.
“As a coach, all I have ever wanted to do was to make a positive impact in the lives of my players and coaches and do so in a way that would benefit them throughout their lives.”
Denison Vice President and Director of Athletics Nan Carney-DeBord is grateful for the leadership Caravana has provided to the program.
“Mike Caravana is a fierce competitor.
He has coached young men throughout his career to compete at the highest level,” said Carney-DeBord. “His coaching record speaks for itself and leaves an impressive legacy. We wish Mike and his family nothing but the best.”
Caravana’s decision to step aside was not taken lightly but he felt the timing was right.
“I am looking forward to starting a new chapter in my life. I have been fortunate to have such a supportive wife in Ashlin, and my three sons, Nick, Sam, and Wyatt. They allowed me to put the necessary time in to have such a wonderful experience at Denison.”
Here’s the bullet point guide to the Mike Caravana Era of Denison Lacrosse.
The Mike Caravana File
28 Seasons (1991-2005, 2009-21)
Career Record: 320-97
Career Winning Percentage: .767
18 NCAA Tournament Appearances
11 NCAA Tournament Quarterfinal Appearances (Elite 8)
3 NCAA Semifinal Appearances (Final 4)
1994 USILA Division III Coach of the Year
2-time IMCLA West Region Coach of the Year
15 NCAC Championships
6 NCAC Tournament Championships (since 2013)
7-time NCAC Coach of the Year
One of 21 men’s lacrosse coaches with over 300 career victories
Charlottesville Chapter Lacrosse Hall of Fame
Ohio Lacrosse Foundation Hall of Fame
Long Island Metro Chapter Lacrosse Hall of Fame
Has Coached…
75 USILA All-Americans
17 USILA Scholar All-Americans
15 NCAC Players of the Year
133 NCAC First-Team selections
3 First-Team USILA All-Americans
1 NCAA Postgraduate Scholarship recipient
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The drills will cover every detail of stick handling, footwork, positioning, checking, defensive holds and creating transition.
Sessions may be shortened to accommodate poor weather. 
morristown summit
Patriot Outreach offers community service opportunities in the lacrosse community and supports community service opportunities of its members.
Our focus is on building the complete student athlete to give them an edge when it comes to taking the necessary steps forward both on and off the lacrosse field.Two-Step Regulation of LAX PANICLE1 Protein Accumulation in Axillary Meristem Formation in Rice
Abstract
Axillary meristem (AM) formation is an important determinant of plant architecture. In rice (Oryza sativa), LAX PANICLE1 (LAX1) function is required for the generation of AM throughout the plant’s lifespan. Here, we show a close relationship between AM initiation and leaf development; specifically, the plastochron 4 (P4) stage of leaf development is crucial for the proliferation of meristematic cells.
Coincident with this, LAX1 expression starts in the axils of leaves at P4 stage. LAX1 mRNA accumulates in two to three layers of cells in the boundary region between the initiating AM and the shoot apical meristem. In lax1 mutants, the proliferation of meristematic cells is initiated but fails to progress into the formation of AM. The difference in sites of LAX1 mRNA expression and its action suggests non-cell-autonomous characteristics of LAX1 function. We found that LAX1 protein is trafficked to AM in a stage- and direction-specific manner. Furthermore, we present evidence that LAX1 protein movement is required for the full function of LAX1. Thus, we propose that LAX1 protein accumulates transiently in the initiating AM at P4 stage by a strict regulation of mRNA expression and a subsequent control of protein trafficking. This two-step regulation is crucial to the establishment of the new AM.
INTRODUCTION
The complex architecture of plants is mainly controlled by the patterns of axis formation. In higher plants, the primary apical-basal axis is established when a shoot apical meristem (SAM) and a root apical meristem are formed during embryogenesis. Starting from this simple bipolar organization, a complex body plan is developed over the lifetime of the plant. The complexity of aerial architecture is created by the repeated generation of new growth axes (McSteen and Leyser, 2005; Schmitz and Theres, 2005). During postembryonic growth, secondary axes of growth are formed as shoot branches. The shoot branches originate from axillary meristems (AMs) that are produced in the axil of the leaves. In vegetative growth, the AMs subsequently produce a few leaf primordia and grow to become axillary buds. Axillary buds are subjected to another level of regulation, becoming dormant or continuing growth, depending on internal and external cues. Therefore, the pattern of vegetative shoot branching is determined not only by the regulation of AM initiation but also by the regulation of bud outgrowth. AMs are also generated in the axils of leaves or bracts in reproductive development and develop into inflorescent branches or flowers. Generation of AMs and determination of their identity are important factors for the establishment of the inflorescence structure.
A number of mutants that exhibit defects in AM formation have been described for several plant species. They are classified into two types according to the characteristics of their phenotypes. One class of mutants, comprising pinhead, revoluta, and cup-shaped cotyledone3 (cuc3), exhibits defects in AM formation in addition to other abnormalities, such as defects in SAM formation during embryogenesis, establishment of leaf polarity, and vascular formation (McConnell and Barton, 1995; Talbert et al., 1995; Otsuga et al., 2001; Hibara et al., 2006; Raman et al., 2008). Therefore, the defective genes in these mutants are probably involved in mechanisms that control general aspects of meristem initiation or function unless AM formation is affected as a secondary consequence of these mutations. The other class of mutants, comprising Arabidopsis thaliana lateral suppressor (las) and regulator of axillary meristem (rax) mutants, tomato (Solanum lycopersicum) lateral suppressor (ls) and blind (bl), maize (Zea mays) barren stalk1 (ba1), and rice (Oryza sativa) monoculm1 (moc1) and lax panicle1 (lax1) show AM-specific defects (Schumacher et al., 1999; Komatsu et al., 2001, 2003; Ritter et al., 2002; Schmitz et al., 2002; Greb et al., 2003; Li et al., 2003; Keller et al., 2006; Müller et al., 2006). These AM-specific mutants can be further divided into two classes: either the mutant shows developmental specific defects or all AMs are affected regardless of the plant’s developmental stage. In ls and las mutants, defects are mainly observed in the vegetative phase. By contrast, bl, ba1, and moc1 exhibit suppression of all types of AM. The molecular basis for this developmental stage-specific regulation of AM formation remains to be elucidated.
Cloning of the genes that are defective in the AM mutants has been reported. LS, LAS, and MOC1 are orthologous genes encoding transcriptional regulators of the GRAS family (Schumacher et al., 1999; Greb et al., 2003; Li et al., 2003). BL and RAX are also orthologous genes that encode R2R3-type Myb transcription factors (Schmitz et al., 2002; Keller et al., 2006; Müller et al., 2006). This shows that AM formation may be regulated by a conserved mechanism in different plant species, including monocots and eudicots. Rice LAX1 and maize ba1 are orthologs and encode basic helix-loop-helix (bHLH) transcription factors (Komatsu et al., 2003; Gallavotti et al., 2004). Although sequence similarity between LAX1 and ba1 is very high, their cognate orthologs have not been found in eudicots, including Arabidopsis. Thus, monocot-specific regulation also operates in the formation of AM. This might reflect the different ways AMs differentiate in different species.
In rice, AMs that arise during the vegetative phase develop as branch shoots called tillers (Hoshikawa, 1989). Upon transition to the reproductive phase, the SAM of each tiller is transformed into an inflorescence meristem and differentiates into an inflorescence, called a panicle. Each inflorescence meristem produces several AMs that develop into rachis branches, and the AMs generated subsequently develop into the next order of rachis branches or lateral spikelets depending upon the time and position of their occurrence. In lax1 mutants, AM formation in the inflorescence is severely suppressed (Komatsu et al., 2001, 2003). LAX1 expression is observed along with AM formation throughout the lifetime of a plant. However, LAX1 expression is not detectable at the stage of SAM formation during embryogenesis, suggesting that LAX1 may be involved in a process that is specific to AM formation.
The mutant phenotype of lax1 implies a possible function of LAX1 in the developing AMs; however, LAX1 mRNA is not detected within the newly initiating AM. Accumulation of LAX1 mRNA is restricted within a few layers of cells at the boundary region between the SAM and the initiating AM (Komatsu et al., 2003). Therefore, it is likely that LAX1 acts in a non-cell-autonomous manner, although its molecular mechanism is unknown. There are many genes that show non-cell-autonomous action as LAX1, but the underlying molecular basis has only been elucidated in a few cases. Cell-to-cell communication through protein trafficking is involved in plant morphogenesis, including root cell patterning and shoot organogenesis (Kurata et al., 2005). In the symplastic intercellular communication, signal molecules, such as maize KNOTTED1 (KN1) and Arabidopsis SHORT ROOT (SHR) proteins, are transported to adjacent cells through plasmodesmata (Nakajima et al., 2001; Kim et al., 2002). LEAFY (LFY), a transcription factor that activates floral homeotic genes, has also been described as an intercellular movement protein (Sessions et al., 2000; Wu et al., 2003). There is evidence suggesting that the activity of the protein trafficking machinery is dependent on cell types and developmental stages allowing protein movement in a highly regulated manner (Kim and Zambryski, 2005). In spite of the progress in our understanding toward the molecular mechanisms controlling protein trafficking, the impact of protein trafficking for its biological function has only been demonstrated in a few studies.
Here, we show that LAX1 function is required for the generation of all types of AMs in both vegetative and reproductive phases. Detailed observation of steps in AM formation indicated a close relationship between AM initiation and leaf development and that plastochron 4 (P4) stage of leaf development is crucial for AM formation. Coincidentally, LAX1 expression starts in the axils of leaves at this stage, and LAX1 protein is trafficked to the AM, which occurs only around the P4 stage. Based on these results, we propose that transient accumulation of LAX1 protein in the AM at P4 stage, which is strictly regulated by mRNA expression and protein trafficking, controls the proliferation of meristematic cells, which is crucial for the full development of AM.
RESULTS
AM Formation in Rice
AMs in grass species develop from the base of the leaf as in most plant species (). However, because they emanate from the shoot side, the meristems are connected to the node in transverse sections. Stages of AM development are indicated as stage of subtending leaves determined according to plastchron number (Pi) system (Sharman, 1942; Hill and Load, 1990). In the axils of leaves at P1 and P2 stages, AMs are not visible, but slight bulging toward the subtending leaf could be observed at P2 stage (, arrowhead). At P3 stage, the lateral bulge of cells that would give rise to the AM became clearly distinguishable (, arrowhead). The number of relatively small and cytoplasmically rich cells increases rapidly between P3 and P4 stages (), and formation of the new AM is completed by P5 stage ().
Axillary Bud Formation during Vegetative Development of Rice.
(A) Longitudinal section of shoot apex stained with safranin-orange G-ferrous tannic acid. The newest leaf is designated as P1, and other leaves are numbered accordingly.
(B) to (F) Transverse sections of the shoot in (A). Arrowheads indicate small bulge that will become the AM.
(G1) to (G17) Localization of OSh2 mRNA that serves as a marker for meristematic cells. The number indicates the distance from the top of the SAM in micrometers. Panels labeled “e” are close-up views of the region delineated by a rectangle in the previous panel. The SAM region is presented as a red circle in (G1e) to (G3e). Arrows in (G4) and (G5e) indicate the margins of P2 leaf. Arrows in (G6e) and (G7e) indicate OSh2 expression underneath the site where P2 leaf margins overlap. Arrowheads in (G7e) and (G8e) indicate the lateral bulge of cells.
(H) A schematic of OSh2 expression in the shoot apex. The region of OSh2 mRNA expression is depicted in red.
Bars = 50 μm.
To investigate the formation of new AMs in more detail, we performed in situ hybridization analysis using rice homeobox 1 (OSh2) mRNA as a marker (). OSh2 is a rice ortholog of KN1 that is expressed in meristematic cells in the SAM (Sentoku et al., 1999). At the top of the shoot, OSh2 is uniformly expressed in the SAM (). At the base of the P1 leaf, OSh2 signal was clearly on the side of the P1 leaf (). Strong OSh2 expression was observed in the SAM at the base of the P2 leaf (). Low, but significant, OSh2 signal was observed in the marginal region of the P1 and P2 leaves where the edge of both sides contact (, arrow). The signal was high underneath the marginal region of the P2 leaf in the lower section (, arrow). Accumulation of newly formed small cells is observed in the axil of the P3 leaf; however, OSh2 mRNA expression could barely be detected in this region in spite of the increasing number of cells in the future AMs ( and arrowheads in 1G7e and 1G8e). Thereafter, OSh2 expression is observed in the axils of the P4 leaves, and the signal was observed in the entire region of the growing AM (). At P5 stage, OSh2 expression is diminished in the incipient leaf primordia and confined to the apical region, suggesting that the new AM has become functional at this stage ().
These observations suggest that cell proliferation starts in the axils of the leaves at P2 stage. Cells that give rise to AMs become distinguishable from neighboring cells at P3 stage. However, these cells do not express OSh2, indicating that they are not yet determined as meristem cells. The fate of the cells as meristem is determined between P3 and P4 stages, and OSh2 expression is detected in the entire region of the future meristem.
We have determined expression patterns of genes that are likely involved in regulating AM formation, including LAX1. Accumulation of LAX1 mRNA was first observed in the axils of the P4 leaves, and the localization of LAX1 mRNA was restricted within a boundary region at the adaxial side of the future meristems (; Komatsu et al., 2003). LAX1 expression remains in the same boundary region but at a lower level at the P5 stage ().
Expression Analysis of LAX1 and Related Genes.
In situ hybridization analysis of LAX1 and related genes mRNA expression in transverse sections of a developing meristem. Column labels indicate the stage of the leaf surrounding the meristem.
(A1) to (A5) LAX mRNA expression.
(B1) to (B5) OSh2 mRNA expression.
(C1) to (C5) Histone h5 mRNA expression.
(D1) to (D5) CDKB2;1 mRNA expression.
(E1) to (E5) MOC1 mRNA expression.
(F1) to (F5) Rice CUC3 mRNA expression.
(G1) to (G5) Rice ra2 mRNA expression.
Bars = 50 μm.
Cell division status during AM formation was examined using Histone h5 and CDKB2;1 as markers. Histone h5 and CDKB2;1 are expressed specifically at the S and G2/M phases of the cell cycle, respectively (Reichheld et al., 1998; Umeda et al., 1999). The two markers showed essentially similar expression patterns in the AM (). In the axils of the P1 leaves, the Histone h5 signal was observed within the SAM (). Despite the fact that future AM cells are barely distinguishable in the axils of the P2 leaves, Histone h5 and CDKB2;1 are not expressed uniformly in these cells. In these cells, Histone h5 and CDKB2;1 expression was localized to the cells adjacent to the future meristem cells but not in the future AM region (). This specific expression pattern became more prominent at P3 stage axils (). Conversely, expression of Histone h5 or CDKB2;1 was frequently observed in proliferating AM cells in the axils of P4 stage leaves ().
Expression of MOC1 and rice CUC3 mRNA appeared to begin in the axils of the P2 leaves (). On the other hand, rice Ra2 mRNA could only be observed from P3 stage (). These genes are known to be expressed at the boundaries between organs (Li et al., 2003; Bortiri et al., 2006; Hibara and Nagato, 2006). We have observed the same expression patterns as previously reported at the boundaries between the shoot and the leaves () and between the shoot and the bud at P3 and P5 stages (). However, at P4 stage, all three genes are expressed in the entire region of the developing AMs ().
LAX1 Is Required for Axillary Bud Formation in Both Vegetative and Reproductive Phases
Previously, we reported that LAX1 and SPA genes are main regulators of AM formation in rice (Komatsu et al., 2003). LAX1 was shown to encode a putative transcription factor containing a bHLH domain, but the molecular nature of SPA had not been determined. The SPA locus was mapped to a region between 81 and 96 centimorgans on chromosome 6 where MOC1 resides (Li et al., 2003). It led us to infer that spa might be an allele of moc1. We determined the nucleotide sequence of a coding region of MOC1 in the spa mutant and it revealed that spa is a new allele of moc1, containing a point mutation that creates a premature termination codon at position Trp-370 (see Supplemental Figure 1 online). Thus, we have renamed this mutant moc1-3. In the double mutant lax1-1 moc1-3, initiation of tillers and vegetative branch shoots is almost completely suppressed, whereas lax1-1 or moc1-3 single mutants are capable of producing tillers. This implies that LAX1 is required for both inflorescence branching and tiller formation. The fact that LAX1 expression is observed during vegetative phase as well as in the reproductive phase corroborates this conclusion. Indeed, we have observed a reduction of the tiller number in lax1-2, which is a strong allele ().
Defects of AM Formation in lax1 Mutants.
(A) and (B) Tiller formation in the wild type and lax1-2 mutant at 30 d after sowing. The lax1-2 mutant has a reduced number of tillers.
(C) and (D) Axillary bud formation in the wild type and lax1-2 mutant. Arrowheads indicate tillers and axillary buds. Yellow arrows indicate the absence of axillary bud in the axils of leaves. The numbers indicate the absolute number of leaves counted from germination. Leaves were removed to show axillary buds.
(E) and (F) Axillary bud formation in the wild type and moc1-3 mutant. The numbers indicate the absolute number of leaves counted from germination. Leaves were removed to show axillary buds.
(G) Tiller bud formation in five lax1 alleles and moc1-3 mutants. The frequency of tiller bud formation in each leaf axil is shown. Twelve plants were examined for each lax1 mutant allele. In this analysis, wild-type plants of all genetic background showed 100% tiller bud formation. The main culm of each plant was examined.
Bars = 10 cm in (A), 4 cm in (B), and 500 μm (C) to (F).
Because shoot branching is controlled by two distinct steps, the formation of axillary buds and bud outgrowth, we have attempted to determine the step that is impaired in lax1-2 and moc1-3 mutants. In wild-type plants, an axillary bud is formed in the axil of each leaf, and the absence of the axillary bud was often observed in the axils of the leaves in lax1-2 and moc1-3, indicating that the decrease of tiller numbers was a result of a failure in the axillary bud formation rather than bud outgrowth (). We have also examined axillary bud formation in five lax1 alleles. In concordance with the severity of the mutations in the LAX1 sequence and inflorescence phenotypes (Komatsu et al., 2003), a variable severity of defects was observed in plants homozygous for the five lax1 alleles (). No defect was detected in lax1-4, the weakest allele, whereas some axillary buds were missing in the moderate alleles, lax1-1 and lax1-5. Absence of axillary buds was more frequent in lax1-2 and lax1-3, which are complete loss-of-function alleles. Notably, formation of axillary buds in the fourth leaf axil was completely abolished in lax1-2 and lax1-3, and fifth and sixth leaf axils showed a tendency to lose axillary buds. This suggests that loss of LAX1 function exerts differential effect on the formation of axillary buds of different leaves and that the fourth leaf axil appears to be affected the most by the mutation. Suppression of axillary bud formation was also observed in moc1-3 (), consistent with a previous report (Li et al., 2003). However, no clear trend of developmental regulation of susceptibility was observed (). Taken together, these results show that LAX1 is involved in a general mechanism that regulates the formation of all axillary buds throughout vegetative and reproductive phases. Considering the additive effects observed in the lax1-1 moc1-3 double mutant and the distinct patterns in the loss of axillary bud formation, the regulatory pathway in which LAX1 functions is likely to be independent from that of MOC1.
LAX1 Function Is Required to Sustain the Processes of AM Formation
To obtain further insight into the role of LAX1 in the control of AM formation, we investigated defects in lax1-2 using OSh2 and Histone h5 as markers. Although impairments of axillary bud formation in lax1 mutants are milder during vegetative phase compared with that observed in reproductive phase, we found that AM formation was completely blocked in the axils of fourth leaves in plants homozygous for strong alleles, such as lax1-2 and lax1-3 (). Therefore, our investigation focused on the development of AMs in the fourth leaf axils and the expression of Histone h5 and OSh2 in the axils of the fourth leaves during their development from P3 to P5 stages. At P3 stage, no differences in the morphology of the plant or expression patterns of the marker genes was observed between the wild type and lax1-2 (). Differences became evident at P4 stage, at which a number of Histone h5–expressing cells are found surrounding the region where new AM is formed in the wild type. By contrast, the region surrounded by Histone h5–expressing cells in lax1-2 is much smaller (). Similarly, OSh2 expression was observed in the entire region of the future AMs in the wild type (), whereas the area and the intensity of OSh2 expression was greatly reduced in lax1-2 (). At P5 stage, the axillary bud was well established in the wild type, but in lax1-2, Histone h5 and OSh2 expression was no longer observed and the meristematic region has disappeared (). Thus, in spite of the complete loss of LAX1 function in the first step of AM formation, shown by the accumulation of meristematic cells in the axils of leaves, is initiated in lax1-2. However, proliferation of meristematic cells is not sustained, leading to the failure of new AM formation. These results indicate that LAX1 function is required for the continuous development, but not the initiation, of AM formation
Expression Pattern of Histone h5 and OSh2 in the Axils of the Fourth Leaf in the Wild Type and lax1-2.
(A) to (D) Expression of Histone h5 ([A] and [B]) and OSh2 ([C] and [D]) in the developing AM in the axil of the forth leaf when the forth leaf corresponded to P3.
(E) to (H) Expression of Histone h5 ([E] and [F]) and OSh2 ([G] and [H]) in the developing AM in the axil of the forth leaf when the forth leaf corresponded to P4.
(I) to (L) Expression of Histone h5 ([I] and [J]) and OSh2 ([K] and [L]) in the developing AM in the axil of the forth leaf when the forth leaf corresponded to P5.
(A), (C), (E), (G), (I), and (K) show the wild type, and (B), (D), (F), (H), (J), and (L) show lax1-2. Bars = 50 μm.
LAX1 Protein Is Trafficked to the Cells in Developing AM
Accumulation of LAX1 mRNA was restricted to two to three layers of cells in the boundary region of the adaxial side of new AMs (Komatsu et al., 2003; ), whereas defects in lax1 are in the developing AMs. This peculiarity suggests that LAX1 acts in a non-cell-autonomous manner to transmit a signal so that cell proliferation continues in the future AM to complete the meristem formation. One possible mechanism underlying this non-cell-autonomous function is that LAX1 protein is trafficked from the boundary region to the future AM cells. To test this hypothesis, we produced transgenic rice plants expressing green fluorescent protein (GFP):LAX1 fusion protein under the control of the LAX1 promoter. Localization of the GFP:LAX1 mRNA was determined to be identical to that of endogenous LAX1 mRNA (), indicating that the LAX1 regulatory region used was sufficient to confer the correct expression pattern.
Trafficking of LAX1 Protein.
(A) In situ localization of GFP:LAX1 mRNA in the developing secondary rachis branch of a transgenic plant. The expression of GFP:LAX1 mRNA was restricted to the boundary region (arrowhead). An antisense GFP probe was used to detect GFP:LAX1.
(B) Confocal image of GFP:LAX1 protein distribution. GFP signals were observed in AM cells in a stage-specific manner (white arrowhead). Yellow arrows indicate the LAX1 protein in the boundary region where the protein was produced. The sample stage is identical to (A). GFP fluorescence is green, and FM4-64–stained plasma membrane is blue.
(C) to (F) Confocal image of transverse sections in the developing axillary buds during vegetative phase. In the P4 stage, GFP:LAX1 protein was observed in the AM cells ([C] and [D]). By contrast, GFP signals were restricted to the boundary region at the early (E) and late (F) P5 stage.
(G) and (H) Confocal image of 3xGFP:LAX1 (G) and GFP:LAX1 (H) fusion proteins in the developing primary rachis branch. GFP signals were limited to the boundary region resulting from the inhibition of cell-to-cell protein trafficking (G). By contrast, GFP:LAX1 protein was shown in AM cells (H).
Bars = 50 μm in (A), (B), (G), and (H) and 100 μm in (C) to (F).
In contrast with the boundary-specific accumulation of GFP:LAX1 mRNA, GFP fluorescence was detected in the AM cells when a small bulge began to appear (, arrowhead). On the other hand, GFP signal was restricted to the boundary region of the adaxial side of the developing AMs at early stage of AM formation when cell proliferation begins and at the late stage when axillary bud formation was completed (, arrows). This suggests that GFP:LAX1 protein accumulation in the meristem is developmentally regulated. In addition to this stage-specific regulation, the intercellular trafficking of LAX1 protein is also directionally regulated. In our analysis, we did not observe expansion of GFP signal toward the stem and SAM. Similar stage- and direction-specific LAX1 trafficking was also observed in AM formation during vegetative phase (). GFP fluorescence was observed in the entire region of the developing AM in the axils of leaves at P4 stage () but restricted to boundary region of adaxial side of new AM at P5 stage (). Only weak GFP florescence was detected in the stem (), which correlates well with the weak expression of LAX1 mRNA in this region (). Taking these results together, we propose that LAX1 protein is trafficked from the boundary region to the developing AMs in a developmental stage-dependent manner.
To confirm that the moving molecules are LAX1 protein and not LAX1 mRNA, we produced transgenic rice plants harboring a 3xGFP:LAX1 fusion gene under the control of the same LAX1 promoter as described above. It was anticipated that intercellular movement of the fusion protein might be inhibited due to the large size of the fusion protein and that the chimeric protein may remain where it is translated (Kim et al., 2005). Indeed, in contrast with the location of 1xGFP:LAX fusion protein, which was detected in the AM, 3xGFP:LAX fusion protein was localized in the boundary region of the developing inflorescence (). Based on these observations, we concluded that the LAX1 protein is synthesized in the boundary region and is trafficked to the initiating AMs. In summary, trafficking of LAX1 protein exhibits three characteristics: first, the LAX1 protein moves in a unidirectional manner toward the AM cells; second, trafficking of LAX1 is developmentally regulated so that LAX1 protein movement occurs only when the cells in the AM are proliferating; third, LAX1 protein trafficking occurs during both vegetative and reproductive phases.
Trafficking of LAX1 Protein Is Necessary for Its Function
To investigate whether the intercellular movement of LAX1 protein is a prerequisite for its function, the 3xGFP:LAX1 chimera gene was introduced into lax1-3, the null mutant allele. In the lax1-3 inflorescence, generation of AMs that give rise to lateral spikelets is completely suppressed, while their subtending bracts are normally formed. Taking advantage of this characteristic, we evaluated the complementation of lax1-3 phenotype as the rate of bracts containing lateral spikelets. Lateral spikelet formation is completely suppressed in the lax1-3. We used 12 and 16 transgenic lines harboring 1xGFP:LAX1 and 3xGFP:LAX1, respectively. All 12 independent 1xGFP:LAX1 plants showed almost full complementation of the lax1-3 mutation. On the other hand, the degree of the complementation varied from very low to full complementation among 16 independent transgenic plants containing 3xGFP:LAX1 (). These results indicate that trafficking of LAX1 protein is required for its full function.
Complementation of lax1 Phenotype by GFP:LAX1 and 3xGFP:LAX1 Fusion Constructs.
(A) and (B) Panicle morphology of representative transgenic plants harboring 1xGFP:LAX1 or 3xGFP:LAX1. These constructs were introduced into the lax1-3 mutant.
(C) Frequency of lateral spikelet formation in the transgenic (1xGFP:LAX1 or 3xGFL:LAX1) and nontransgenic (WT) plants. The ratio of lateral spikelets relative to the number of bracts, modified leaves, in a panicle is shown. 1xGFP:LAX1 or 3xGFP:LAX1 genes under the control of the LAX1 regulatory region were introduced into lax1-3 mutants. The first regenerated T1 plants were used for the analysis.
To investigate if the 3xGFP:LAX1 fusion protein prevents LAX1 from functioning, we have used pINDEX two-component inducible system (Ouwerkerk et al., 2001) to produce a transgene encoding 3xGFP:LAX1 fusions that are inducible by dexamethasone treatment. We previously reported that rice DOF works downstream of LAX1 and its transcription is upregulated upon the induction of LAX1 expression (Furutani et al., 2006). Introduction of pGFP:LAX1-INDEX and p3xGFP:LAX1-INDEX into lax1-3 calli resulted in a similar level of DOF upregulation upon dexamethasone treatment (see Supplemental Figure 2 online), indicating that the molecular function of LAX1 protein as a transcription factor was not affected by fusion with 3xGFP protein. This data corroborates the conclusion that the failure of full complementation of lax1-3 phenotype was due to the lack of movement of the 3xGFP:LAX1 fusion protein.
DISCUSSION
AM Formation in Rice
Two possible theories have been proposed for the formation of AM: (1) that it is generated by de novo occurrence of new meristematic cells or (2) that it results from the detachment of existing meristem cells in the SAM (Steeves and Sussex, 1989). Recent studies using molecular markers have shown that, in Arabidopsis, AM formation is initiated from a small number of meristem cells that are derived from the SAM and remain in the center of the leaf axil, supporting the detached-meristem theory (Long and Barton, 2000; Greb et al., 2003). However, the data presented in this study suggest that AM formation may take place in a different way in rice. In the case of rice, cell proliferation for the future AM occurs in the axils of leaves at around P2 stage. However, judging from the expression of OSh2, a marker widely used for the indeterminate cells, these cells are not yet meristematic even at P3 stage. Subsequently, all or most cells in the cluster acquired meristematic fate by P4 stage. Therefore, we did not detect cells expressing OSh2 in the leaf axil until P4 stage, which is in favor of de novo AM synthesis theory. Moreover, it seemed that the proliferation of cells to form a future AM started prior to the onset of OSh2 expression. This also supports the notion that rice AM is not generated as a result of proliferation from a small number of meristematic cells, rather, of an acquisition of indeterminate cell fate by existing cells. We cannot rule out the possibility that a few cells weakly expressing OSh2 are preserved and are responsible for AM initiation. If this is the case, rapid propagation of the meristematic cells as well as the enhancement of OSh2 expression level should be postulated to explain the OSh2 signal in the axils of the leaves at P4 stage. Another possible explanation may be that OSh2 is not a plausible marker for the early stage of meristem formation and an earlier marker could show the presence of meristem cells in the axils of P1 to P3 leaves. However, we consider this unlikely because OSh2 is generally used as an early marker for SAM formation in embryogenesis (Sentoku et al., 1999). Furthermore, even if this is the case, an abrupt onset of OSh2 expression in the entire region of the future AM at P4 stage indicates that a substantial change of cell fate occurs from P3 to P4 stage.
The differences in the mechanism of AM formation may reflect the anatomical divergence of AM origination among species. In grass species, leaves develop to enclose the SAM and the AM in the n-th leaf axil initiates immediately below the site where leaf margins of n+1-th leaf fuse (). Thus, AM formation in rice is closely related with leaf differentiation. Analysis of AM initiation in the context of interaction with leaf development may be needed for further understanding of AM formation in grass species. Absence of LAX1/ba1 genes in eudicots is an additional strong indication of a grass-specific mechanisms controlling AM formation.
LAX1 Functions throughout Development
We propose that formation of rice tiller buds could serve as a model for the analysis of the AM formation process because the progression of each step is relatively slow and constant, and it interconnects well with the development of its subtending leaf.
In rice, three different lateral branches, tillers, panicle branches, and spikelets, are produced. Here, we showed that LAX1 function is required for AM formation throughout the lifetime of rice, and the formation of all three types of branches are affected in lax1 mutants, which correlates with the fact that LAX1 is expressed in all AMs. The ba1 gene, maize LAX1 ortholog, is also required for the formation of AMs in both vegetative and reproductive phases (Ritter et al., 2002; Gallavotti et al., 2004). Therefore, LAX1 appears to be involved in a common regulatory machinery that is essential for AM formation. However, even in the complete loss-of-function mutants, such as lax1-2, in which an entire region of the LAX1 locus is deleted from the genome (Komatsu et al., 2003), some AMs are still generated normally and grow as branches. This suggests that other factors are also required and may work with LAX1 redundantly to support the formation of AMs. The phenotype of the lax1 moc1 double mutant is more sever than either single mutant, and AM formation is completely abolished. In addition, developmental regulation of these genes appears to be different. We propose that MOC1, a rice ortholog of LAS/LS, works independently from LAX1 for the regulation of AM formation. Functions of the recently identified regulators of AM formation, RAX and CUC genes, are yet to be determined in rice (Hibara et al., 2006; Keller et al., 2006; Müller et al., 2006; Raman et al., 2008).
Molecular Basis of Non-Cell-Autonomous Function of LAX1
We showed that LAX1 is one of the non-cell-autonomous proteins (NCAPs). To date, many plant NCAPs have been described (Haywood et al., 2002; Kim, 2005). There are at least two modes of protein trafficking between cells (i.e., targeted trafficking or nontargeted diffusion) (Kim, 2005; Kurata et al., 2005). Targeted trafficking involves specific interactions between signal motifs in NCAPs and plasmodesmatal components, accompanied by plasmodesmata size exclusion limit (SEL) increase. KN1, SHR, and viral movement proteins are known NCAPs that move between cells using targeted trafficking (Carrington et al., 1996; Nakajima et al., 2001; Kim et al., 2002). By contrast, nontargeted trafficking is diffusion based and depends upon the endogenous plasmodesmata SEL. LFY and GFP are examples of this type of NCAP. We infer that LAX1 is also trafficked via a nontargeted mechanism because the molecular weight of LAX1 protein (23 kD) is similar to that of GFP (27 kD) and smaller than that of LFY (47 kD). Moreover, suppression of intercellular movement of LAX1 by adding three molecules of GFP to make 3xGFP:LAX1 fusion protein (104 kD) supported this assumption. The dependence of intercellular movement on molecular size is one of the prominent characteristics of nontargeted movement (Zambryski and Crawford, 2000). It has been reported that the trafficking ability of NCAPs by diffusion correlates with cytosolic localization (Crawford and Zambryski, 2000; Wu et al., 2003). We showed that a significant portion of LAX1 protein was present in the cytoplasm, in addition to its predominant localization in the nucleus (see Supplemental Figure 3 online). However, it is still unclear whether a signal sequence, which is required for targeted movement, exists in the LAX1 protein.
We demonstrated that the trafficking of LAX1 protein is stage specific. The SEL of plasmodesmata changes temporally and spatially to allow the passage of macromolecules (Zambryski and Crawford, 2000). In general, young tissues have greater symplastic connectivity via plasmodesmata than differentiated tissues, thereby allowing more frequent cell-to-cell communication (Gisel et al., 1999). Taking this into consideration, we infer that intercellular movement of LAX1 protein might occur only when the cell division activity is high in the AM at around P4 stage. Once the axillary bud is established, movement of LAX1 may be blocked due to the reduction of the SEL. In addition to the stage specificity of trafficking, LAX1 protein moves only from boundary region toward the AM, but never toward the SAM. A possible explanation for this unidirectional movement is a symplasmic isolation mechanism, where plasmodesmata are closed or restricted at a boundary region (Heinlein, 2002; Ding et al., 2003). We postulate that the AM is symplasmically isolated from the SAM; thus, LAX1 protein cannot move across the boundary region, but only toward the AM.
Two-Step Control of Spatial and Temporal Distribution of LAX1 Activity and Its Implication in Its Function
We have revealed a two-step regulation of spatial and temporal distribution of LAX1 activity, which is essential for its full function. LAX1 mRNA is specifically expressed in the boundary region between the initiating AM and the SAM; subsequently, LAX1 protein is trafficked toward the AM. Therefore, spatial distribution of LAX1 protein is regulated by two steps: cell type–specific mRNA expression and protein trafficking (). Moreover, temporal activity of LAX1 is also controlled by a similar two-step regulation. LAX1 mRNA is expressed from P4 to later stages, while the LAX1 protein movement is restricted to P4 stage. In this way, stage- and cell type–specific LAX1 mRNA expression is further refined by stage- and direction-specific LAX1 protein movement. What could be the functional implication of this type of two-step regulation? Considering that LAX1 protein is targeted to the site where meristematic cells multiply at P4 stage and when cells in the future AM proliferate actively, it is highly possible that the strict control of LAX1 protein is a prerequisite for the strict control of meristematic cell proliferation in the future AM. In concordance, constitutive expression of LAX1 seriously perturbed normal development of rice (Komatsu et al., 2003).
Model of LAX1 Function in the AM Formation.
LAX1 mRNA is specifically expressed in the boundary region when the subtending leaf is at P4 and P5 stages. At P4 stage, LAX1 protein synthesized in the boundary region moves toward the future AM and enhances cell proliferation. In the absence of LAX1 function, the cell proliferation does not take place. As a consequence, the AM is not established. LAX1 mRNA accumulation is shown in blue, and the LAX1 protein distribution is shown in green.
However, in possible contradiction with our model, partial rescue of the lax1-3 phenotype by 3xGFP:LAX1, which is expected to not move, was observed. As shown in , the degree of complementation varied in 3xGFP:LAX1 lines, and only a few lines showed significant levels of complementation. This is in marked contrast with the results that all 1xGFP:LAX1 plants showed almost complete rescue of the lax1 phenotype. The difference between 3xGFP:LAX1 and 1xGFP:LAX1 in their ability to complement lax1 phenotype appears to be related to the location of the LAX1 proteins. We showed that, in spite of LAX1 trafficking toward the growing AM primordia, some LAX1 proteins remain in the boundary, the site of LAX1 protein synthesis. Thus, trafficking of LAX1 results in the expansion of the region where LAX1 proteins are present but does not lead to the complete delocalization of LAX1. We infer that the partial complementation conferred by 3xGFP:LAX1 indicates the functionality of the remaining LAX1 at the site of synthesis. The difference in the degree of complementation between 1xGFP:LAX1 and 3xGFP:LAX1 raises the possibility that LAX1 in the boundary and in the AM operates differently for the establishment of AM. Apparently, LAX1 proteins that moved to the AM have a greater effect than those remaining in the boundary. Determination of LAX1’s functionality in different locations would require the elucidation of the molecular mechanisms by which LAX1 operates. LAX1 encodes a noncanonical bHLH transcription factor, which is predicted not to bind DNA. Therefore, trafficked LAX1 protein probably interacts with its partner protein(s) to regulate downstream targets. It will be of great interest to isolate the partner of LAX1 as well as downstream target gene(s).
Control of AM Formation by Auxin
In the mutants of the ba1 gene, an ortholog of LAX1, AM formation is severely suppressed, while subtending bracts are produced normally (Ritter et al., 2002; Gallavotti et al., 2004). Treatment with N-1-naphtylphthalamic acid, an inhibitor of auxin transport, abolished bracts in the ba1 mutant, implying that auxin transport is required for bract formation in the maize inflorescence (Gallavotti et al., 2008b). When wild-type maize plants were treated with N-1-naphtylphthalamic acid, generation of AMs as well as bract formation was severely suppressed (Wu and McSteen, 2007; Gallavotti et al., 2008b). Therefore, it is likely that auxin transport is required for bract formation, which, in turn, becomes a prerequisite for AM initiation. Concordantly, mutation in bif2 and spi1, carrying defects in the control of auxin transport and biosynthesis, respectively, exhibited the barren inflorescence that lacks both bracts and AM (McSteen and Hake 2001; McSteen et al., 2007; Gallavotti et al., 2008a).
Auxin maxima that are generated at the site of AM initiation in the wild-type inflorescence were not observed in ba1 (Gallavotti et al., 2008). Considering that auxin transport is required for organ initiation (Reinhardt et al., 2003) and bracts are formed normally in the ba1 inflorescence, auxin transport machinery is not impaired in the ba1. Therefore, the absence of auxin maxima in the ba1 inflorescence might be a secondary effect of the failure in AM formation. An alternative explanation is that auxin maxima are established for bract (leaf) initiation and AM formation independently, and ba1/LAX1 controls auxin transport specifically at the site of AM initiation.
Recently, physical interaction between BA1 and BIF2 was demonstrated (Skirpan et al., 2008). Bif2 is a coortholog of Arabidopsis PINOID that controls auxin transport (McSteen et al., 2007). Considering the distinct mutant phenotypes of bif2 and ba1, suppression of bract initiation in bif2, and AM formation in ba1, this finding was surprising and its functional relevance should be tested. However, if this is the case, an interesting scenario emerges: rice PID, a bif2 ortholog in rice that is expressed in the AM (Morita and Kyozuka, 2007), could be one of the candidates that works with LAX1 in the growing AM and that plays a role in sustaining meristem cell proliferation.
METHODS
Plant Material and Growth Conditions
The lax1 mutant alleles and spa/moc1-3 mutant have been described (Komatsu et al., 2003). For measurement of axillary buds, the seeds of lax1 mutants and their wild type were grown in the glasshouse under natural light conditions.
Construction of Vector and Rice Transformation
Construction of vectors is described in the Supplemental Methods online. Rice (Oryza sativa) transformation was performed as described by Hiei et al. (1994).
Histological Analysis
For histological analysis, shoot apices were fixed with FAA (5% formalin, 5% acetic acid, 45% ethanol, and 45% water) and embedded in PARAPLAST PLUS (McCormick). Transverse and longitudinal sections of 12-μm thickness were stained as follows: 2% zinc chloride solution for 1 min, 0.004% safranin and 0.1% ethanol solution for 5 min, 2% orange G and 5%,tannic acid solution for 1 min, 5% tannic acid for 5 min, rinsed with water, and finally 1% ferric ammonium alum solution for 2 min.
In Situ Hybridization
In situ hybridization was performed as described by Kouchi et al. (1995). Antisense digoxigenin (DIG)-labeled LAX1, OSh2, and Histone h5 probes were generated as described previously (Itoh et al., 1998; Komatsu et al., 2003). Partial cDNAs of MOC1 and rice CUC3 were PCR amplified using following primer sets: for MOC1, 5′-GATGATGTCCTGTCAAGGAAG-3′ and 5′-GCAGTAGTGATCGATCGATCA-3′; and for Os CUC3, 5′-TGCCGCCGGGGTTCAGGTT-3′ and 5′-TCATGCGCCCCTGGGCACT-3′. These PCR fragments were cloned into pGEM-T Easy and linearized with appropriate restriction enzymes. To make antisense probes, in vitro transcription was performed using the linearized plasmids as templates, incorporating DIG-UTP. An antisense probe for rice Ra2 was prepared using full-length cDNAs as a template.
Analysis of GFP Fluorescence under Confocal Microscopy
Hygromycin-selected T1 generation of transgenic plants were grown in a growth chamber. Shoot apices were cut by hand and mounted in 50 μg/mL FM 4-64 (Invitrogen) aqueous solution to stain plasma membrane. Confocal images were obtained using an FV1000 confocal laser scanning microscope (Olympus).
Accession Numbers
Sequence data from this article can be found in the GenBank/EMBL/DDBJ databases under the following accession numbers: {“type”:”entrez-nucleotide”,”attrs”:{“text”:”AB115668″,”term_id”:”33342171″,”term_text”:”AB115668″}}AB115668 (LAX1), {“type”:”entrez-nucleotide”,”attrs”:{“text”:”D16507″,”term_id”:”452235″,”term_text”:”D16507″}}D16507 (OSh2), {“type”:”entrez-nucleotide”,”attrs”:{“text”:”AY242058″,”term_id”:”30142080″,”term_text”:”AY242058″}}AY242058 (MOC1), {“type”:”entrez-nucleotide”,”attrs”:{“text”:”AK059682″,”term_id”:”32969700″,”term_text”:”AK059682″}}AK059682 (rice CDKB2;1), {“type”:”entrez-nucleotide”,”attrs”:{“text”:”AK105527″,”term_id”:”32990736″,”term_text”:”AK105527″}}AK105527 (rice Ra2), and {“type”:”entrez-nucleotide”,”attrs”:{“text”:”NM_001068747″,”term_id”:”297608829″,”term_text”:”NM_001068747″}}NM_001068747 (rice CUC3).
Supplemental Data
The following materials are available in the online version of this article.
Supplemental Figure 1. Schematic Representation of moc1-3 Mutation.
Supplemental Figure 2. Investigation of 3xGFP:LAX Protein Activity.
Supplemental Figure 3. Subcellular Localization of LAX1 Protein.
Supplemental Methods. Construction of Vectors, RT-PCR Analysis, and Analysis of Subcellular Localization of LAX1 Protein.
90,000 Why do indoor plants wither? Causes. What to do? – Botanichka.ru
The main symptom of any problems with indoor plants, first of all, is their depressed appearance. Loss of freshness, attractiveness, slight changes in the color of the leaves, which appear more and more obvious – an indication of lack of care or the appearance of various diseases.
A withering houseplant. © KenTannenbaum
Contents:
Why and how do indoor plants wither?
A lethargic or depressed look is a symptom that is too general and vague, it is difficult to determine the cause that led to the appearance of problems.Plants wither, wither, lose their healthy appearance in different ways, very often the depressed state proceeds so individually that even in identical plants it is diagnosed with difficulty and in different ways. Difficulties in identifying the symptoms cause the corresponding difficulties and the search for means to improve the condition of the green pet.
Withering as a plant condition should not be confused with the natural process of partial dropping, wilting and yellowing of foliage . For many indoor plants, getting rid of old leaves is part of the natural development cycle.These include deciduous bushes and trees, plants with a full dormant period, such as anredera, cyclamen, tuberous begonias, caladium, etc. Partial leaf shedding is a normal process for jasmine, fuchsia and camellias, sinningia and coleria, their yellowed sluggish leaves do not indicate on the general depressed state of the plant. As well as the sluggish bracts of poinsettia or the natural process of dying off the lower leaves as the young greenery grows in the palm trees. Before looking for problems and difficulties, it is worth studying the characteristics of the plant and separating natural wilting from the general oppressed, lethargic species due to problems with conditions, care, diseases or pests.
If it seems to you that your plant is withering, you should take a closer look and determine exactly how this happens :
- The initial symptoms of problems are lethargic, slightly drooping leaves that have lost their usual elasticity, slowed growth and a general faded, stale appearance. It is easy to confuse them with signs of a lack of moisture, but, as a rule, we are talking about wilting only when, with sufficient moisture in the substrate, a depressed state remains. The first symptoms will not allow you to identify the cause that led to the problems, but they will indicate that you need to look closely at the plant.
- Nonspecific wilting. It occurs if wilting increases, the plant loses its decorative effect, but other symptoms do not appear in the form of a change in color (paleness, yellowing or browning) or obvious signs of infection with diseases that could cause a depression.
- A general discoloration, slight, barely noticeable discoloration or fading of greenery and shoots is a secondary symptom indicating serious problems and what caused them.The changing color is associated with the destruction of chlorophyll and impaired photosynthesis, metabolic problems. The changing color of greenery, along with a general sluggish appearance, directly indicates problems. There are three types of discoloration of depressed and withering plants:
- blanching of leaves, initial or slight chlorosis;
- yellowing of leaves;
- russeting of aboveground parts.
Identifying symptoms is the first step in diagnosing and fixing a problem.For sluggish-looking houseplants, you need to analyze all the factors of care and compare them with the preferences of the plant in order to immediately find the main reasons that could cause the wilting process. But sometimes a simple analysis of the conditions or irrigation regime with additional fertilizing is not enough.
A depressed look in indoor plants can be provoked by:
- Wrong watering
- Insufficient feeding
- Unfavorable indoor climate
- Pest infestation
- Spread of fungal and viral diseases
Simple correction of care and conditions most often allows you to cope with the primary symptoms and prevent further deterioration of the pet’s condition.But if we are talking about nonspecific wilting or changing color, then the establishment of only care is indispensable. This is just the first step towards identifying and fixing the problem.
Blanching of leaves and wilting of a houseplant. © rickrack54
Slight blanching of leaves: symptoms, causes and methods of control
A slight degree of chlorosis or general blanching of leaves is clearly visible against the background of a general depression. Sluggish and discolored leaves are striking against the backdrop of healthy indoor plants, the appearance becomes more nondescript and painful, the plant resembles a pale version of its former beauty.Dull color, discoloration, blanching of only part of the leaves or a specific zone of limp leaves indicate various health problems of the plant.
| Symptoms of Problem | Reason for wilting |
|---|---|
| Flabby leaves that have lost turgor are combined with a dull color, without timely measures taken, the leaves sag more and more and brown spots grow on them, in a neglected state some of the leaves fall off | Excess watering and spread of root rot |
| Blanching or complete discoloration (uniform over the entire crown or expressed to varying degrees), the appearance of watery, dark areas, a sluggish drooping appearance | Hypothermia or frostbite |
| The lethargic appearance is combined with discoloration of both leaves and shoots; gradually depressed state is supplemented by stretching the plant and shrinking leaves | Low illumination |
| A sluggish-looking plant initially discolours, and then the leaves begin to turn brown and die off | Thrips Defeat |
| The plant seems pale and lethargic, it is in a neglected state, the leaves turn unevenly along the crown, turn yellow and dry out | Spider mite infestation |
| Sluggish leaves do not turn pale, but the plant is deformed, dwarfism develops and signs of growth problems appear | Defeat by nematodes |
| Slow discoloration starting from the bottom of the crown and from the veins of the leaves, which is gradually replaced by yellowing and browning of the leaves | Lack of nitrogen |
| Blanching of the reverse side of the leaf plates between the veins, gradually covering the entire leaf and turning into purple erosion along the edge of the leaves | Lack of magnesium |
| Blanching of the lower leaves, sharply turning into brown | Potassium deficiency |
| Blanching of the upper leaves, sharply turning into brown when neglected | Lack of iron |
| Blanching of the base of old leaves and tips of young leaves, which gradually turns into leaf death | Boron shortage |
| Blanching of the tissues of the leaf plate only between veins with gradual yellowing and repainting in orange | Lack of manganese |
| Pale spots on leaf plates, which, when neglected, turn into specks from rusty spots, and repainting in a bronze shade | Zinc deficiency |
It is especially worth choosing a method of treating a plant when it is damaged by pests and diseases.It is always best to try to deal with this problem by washing, correcting care and biological methods before applying chemicals. But if the damage is serious or the situation is getting worse, insecticidal drugs are the only chance to save the plant.
Methods of dealing with problems directly related to their cause:
- Correction of watering helps only at the initial stage of wilting. If the leaves turn pale, then the only measure to combat rot and dampness, which led to wilting, is an emergency transplant with the removal of damaged tissue.The same method – albeit risky, but a chance to get rid of the consequences of soil contamination with nematodes.
- Raising the temperature and removing damaged leaves and shoots will help save a plant that has suffered from hypothermia.
- Supplementary lighting or a light sanatorium will help solve the problem of insufficient illumination.
- Infusions of onion and garlic or insecticides (celinone, karbofos, decimus, potassium soap) will help to cope with the defeat of thrips.
- If the depressed appearance of a pale plant is caused by a spider mite, then simply increasing the humidity to combat the problem will not be enough.Moisture therapy for tropical plants, the use of decoctions, and infusions (tops of tomatoes, bitter wormwood, chamomile, dandelion) and insecticides (from malophos to pyrethrum, carbolineum, acaricide, potassium soap) will help to cope with various stages of spider mite infestation.
- The introduction of highly specialized and monofertilizers (nitrogen, potassium, boric, etc.) helps to cope with pallor and wilting caused by a lack of certain macro- and microelements.
If the depressed state of the plant is associated with problems that cannot be solved, for example, severe frostbite or decay of the base of the shoots, then it may not be possible to save the plant.Cutting and rooting cuttings will help grow a new plant to replace the old one.
Withering of a houseplant
Causes and elimination of yellowing of leaves
Drooping, lethargic leaves and an unhealthy appearance of a plant with yellowing are not combined so often, but it is also much easier to diagnose the cause of the depressed state of the plant than in the case of blanching, the obvious symptoms of which usually appear at serious stages of the lesion. In all cases, when the yellowing of the leaves was preceded by their blanching, it is necessary to deal with the problems of blanching.But if wilting is accompanied only by yellowing, then we are talking about several other problems – an excess, not a lack of fertilizers, insufficient watering and diseases.
| Symptoms of Problem | Reason for wilting |
|---|---|
| Sluggish leaves gradually turn yellow at the tips, the process spreads further, in a neglected state the tips of the leaves turn brown and dry, and the plant stops growing and stops flowering | Poor watering, frequent drought |
| Before yellowing, the leaves not only wither, lose turgor, but also curl | Damage to aphids |
| On dull leaves, yellowing gradually spreads, starting from the tips, after which weeping spots on greenery and stems are noticeable, indicating a problem | Defeat with wet bacterial rot |
| The foliage gradually turns yellow more and more, and the smallest black dots appear on the underside, somewhat reminiscent of mold | Dry rot damage |
| In a sluggish-looking plant, the stem and the underside of the leaves turn yellow, especially in the area adjacent to the root collar; when neglected, the root collar turns black | Fusarium disease |
| Yellowing and sluggish appearance is characteristic mainly of the lower leaves.Flowering is gradually disrupted – color changes, peduncles are shortened | Excess potassium |
| The oldest leaves of a dull plant turn yellow, there are signs of premature or accelerated aging | Excess phosphorus |
If the yellowing of a dull plant is caused by insufficient watering , then the problem is solved by correcting watering and dividing the usual procedures into frequent, but not abundant ones. Usually, water for irrigation is applied at intervals of several hours a couple of times or saturated with an earthen lump by immersion in water, further maintaining the moisture level that is suitable for a given plant.
Infection of plants with aphids always manifests itself not only in visible traces of insects, but also in a depressed state and difficult recovery. You need to fight this stubborn pest with diligence: even if you managed to cope with the problem, unnecessary preventive treatment will not hurt. After all, one female insect is enough to start everything again. To combat aphids, you can use scenting and washing, fuming, biological agents (decoctions and infusions of dandelion, potato and tomato tops, pyrethrum and chamomile, nettle, wormwood, yarrow, tobacco) or insecticides – from simple household and potassium soap to celinone, detoil , karbofos and modern drugs.
Wet bacterial rot is most often associated with the purchase of plants from unverified sellers, private gardeners and florists using contaminated soil. You can deal with the problem by treating the problem with copper-containing preparations (copper-soap mixture, Bordeaux liquid, copper oxychloride) or with chemicals such as nitrafen, oxychom, phthalan and ferrous sulfate. But if you ignore the yellowing and lethargy, you will most likely not be able to cope with the disease.
With dry rot it is better to immediately fight with the use of insecticides and biological preparations. Both the aboveground parts of plants and the soil need to be processed. You can use infusions of marigold, mullein or wood ash, horsetail decoction, Bordeaux liquid and any systemic fungicides of a wide spectrum of action.
Fusarium can also be fought with biological agents (infusions and decoctions of horsetail, mullein, ash, marigolds) and various fungicides (from copper oxychloride, Bordeaux liquid, ferrous sulfate and phthalan to systemic drugs).This is a very dangerous disease, the delay in the treatment of which can cause the death of the plant.
With an excess of any fertilizers can only be dealt with by stopping feeding, adjusting the fertilizer composition and changing the substrate if its reaction and composition are not suitable for the plant.
A withering houseplant. © plantz
Symptoms and problems of leaf browning
Browning, drying out of individual parts of sluggish plants are most often symptoms of serious stages of damage, following the usual signs – blanching or yellowing.But there are two exceptions to this rule: in indoor plants, the tips of the leaves turn brown and dry out, not only with a lack of nutrients or other problems, but also with too bright lighting and high air temperatures. If browning appears without discoloration or yellowing, it is very easy to identify the problem.
Too bright lighting, exposure to direct sunlight and watering in the midday sun, inadequacy of the sunny location to the habits and requirements of plants often leads to the fact that the leaves, and sometimes young shoots, wilted.Withering, with the regular repetition of these conditions, also leads to the appearance of brown spots on the leaves, traces of sunburn and the beginning of drying out. There is only one method to deal with such browning – immediate correction of conditions. If you move your pet to a less intensely lit place in time, you can avoid a significant loss of decorativeness. If large areas of leaves are brown, then they will have to be cut off or completely removed.
The edges of leaves that have lost their turgor will turn brown, and when kept in heat in those plants that prefer cool conditions.This reaction is typical for almost all indoor plants that require careful selection of conditions. If high temperatures are combined with dry air, then browning spreads faster, and sluggish leaves very poorly restore their usual elasticity. To deal with the problem of browning, you need to immediately correct the conditions – by moving the plant to a more suitable temperature regime.
How to deal with non-specific wilting?
If the houseplant looks lethargic, but all possible symptoms accompanied by changes in the color of the leaves are excluded, we are talking about non-specific wilting.And it is always caused, without exception, by mistakes in care and selection of conditions.
Lead to non-specific wilting:
1. Lack of top dressing , irregular or poor dressing that does not meet the requirements of the plant. They cause depression in the same way as the lack of transplants, the use of poor-quality poor soil. In this case, the plants lack both macro- and microelements, it looks sluggish, growth almost stops.
Solution to the problem: transplanting into fresh soil and regular feeding
2. Drafts , contraindicated for most indoor plants, with the exception of very hardy species and those that have moved into summer rooms, and garden stars.
Solution to the problem: it is enough to put the plant in a protected place and isolate it from active air currents.
3. Inconsistency of temperature and lighting with plant habits . Shading or, conversely, too bright lighting, low or high temperatures can lead to wilting without other obvious signs.
Solution to the problem: it is enough to study the preferences of a particular plant species and put it in a place corresponding to them.
4. Constant wetting of the base of shoots or leaves , leading to the development of rot, especially in the break of the stem.
Solution to the problem: only careful watering will not help, you need to treat the plant with systemic fungicides of a wide spectrum of action.
5. Hypothermia or frostbite of the roots , resulting from exposing the plant in autumn and winter to a cold floor or window sill, transportation in cold weather with crown protection, but without pot protection.
Solution to the problem: emergency transplant with examination of the root ball and removal of damaged parts of the roots.
Non-specific wilting is common among bulbous plants, triggered by the purchase of contaminated planting material or the use of contaminated soil. Blue mold and sclerocial rot, which can be diagnosed by bluish bloom and light accumulations of fungus, respectively, require the use of systemic fungicides. If the bulbs look sluggish, there are no signs of other problems (or possible reasons for wilting associated with leaving), then you should immediately carefully dig them out of the soil and subject the bulbs to an inspection in order to more accurately determine the problem.
How to revive an orchid with sluggish leaves?
Dead orchid. Sluggish leaves. How to get turgor back? Read about resuscitating a sluggish orchid at home. 2 simple solutions for treatment.
How to revive an orchid with sluggish leaves?
Sluggish, rotten, broken, yellowed leaves – there are 2 proven boards for all of the listed manifestations of orchid malaise. First, let’s figure out where to start help.
Orchid resuscitation: step # 1
Look for rotten roots.Cut and sprinkle with fungicide to prevent pathogens from spreading to healthy tissue.
Take the same steps for the leaves and peduncle. Do not pick or cut off dull or yellow leaves – leave them even if there are no roots. The orchid will increase them in the process of resuscitation and gradually strengthen growth.
How to reanimate an orchid: step number 2
Prepare one of the proposed recipes and process limp leaves and growth points according to the instructions. Restore, following the criteria suggested in the third part, so that the orchid quickly regains its shape.
Resuscitation of orchid leaves with succinic acid: 1 tablet per 1 liter of water.
How to properly dilute succinic acid? The drug is available in the pharmacy and is dispensed in tablets or fine crystalline powder. For a solution based on the powder version, draw on the tip of a knife and dilute in 0.7 l of water.
Dissolve a tablet of succinic acid in 1 liter of water. At the dosage suggested by the manufacturer, for example, 1 gram, use the same volume of water as in the tablet form.
Reanimation of orchids with cytokinin paste: 1 mg (bead on a toothpick) locally.
Cytokinin paste – a preparation not only for the restoration of orchids, but also a stimulating ointment for building up children, closing wounds, stimulating renewal. A useful thing, to put it simply. Read more here: how to revive the phalaenopsis orchid?
How to use the products:
- Moisten a cotton pad in a solution of succinic acid and wipe the damaged area several times a day.
- Apply a pea of cytokinin paste to the damaged area, flaccid area, at the base of the branch.
Wrapping after resuscitation of leaves
Use microelements or complex fertilizers a few months after the restoration of the orchid. For flowering plants, dilute 2 times with water and add in small portions. For orchids – according to the instructions on the preparation.
How to feed an orchid with sluggish leaves:
- Bona Forte;
- Pocon;
- “Dr. Foley, Orchid”;
Caring for the orchid for restoration
To restore, dry the soil mixture with the orchid, put it in a shaded place in the house, humidify the air.Room temperature and without sudden changes. Increase watering frequency gradually. Fertilize in 2 months. Inspect the orchid and check the condition every week. Change the approach and amount of watering depending on the state of the orchid.
Orchid leaf renewal course : from 1 week to 1.5 months. We continue regenerating care: why is phalaenopsis not blooming?
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Phalaenopsis orchid sluggish leaves
Sometimes it happens that the leaves of an orchid wither.Before doing anything it is important to establish the reason why it happened. About why phalaenopsis loses turgor, and what to do with it, we will tell you in our article.
Why Phalaenopsis leaves lose their elasticity
There can be several reasons why orchid leaves wither:
- Direct sunlight causes plants to overheat. As a result, rapid evaporation of moisture begins and the leaves dry out. In addition, the substrate gets very hot.Excessive heat is harmful to the orchid. Although Phalaenopsis is a light-loving plant, it prefers diffused light and high humidity. From the heat, the leaf blades wither due to the loss of moisture in the tissues.
- If the orchid wilted in winter, it could be due to the cold . This happens when the temperature is below +15 0 . For phalaenopsis, leaf damage at a low temperature for it is considered frostbite. This means that the leaf blades are lost irretrievably, they cannot be restored in any way.It remains only to cut to living tissue.
- Phalaenopsis are afraid of drafts. A sharp gust of air masses weakens the plants. They are sensitive to adverse influences, loss of turgor (leaf elasticity).
- Violation of the watering regime affects the general condition of the orchids. They do not like waterlogging or drought. Both can cause wilting.
- Root problems will definitely affect the leaves. The root system is bright green with a silvery bloom, indicating the good health of the plant.As soon as spots or drying islands appear on the roots, it means that problems have arisen. Moreover, not only with roots, but also with the ground part.
- Also , leaf wilting can cause diseases and pests of orchids.
Once the reason is established, you can start saving the orchid.
In a healthy orchid, the leaves are dense and point upward and outward. If the plates are hanging, then something is wrong with the plant, and an urgent need to look for the cause of wilting.
What to do when Phalaenopsis leaves withered
Change lighting
If the original location for the phalaenopsis was unsuccessful, it needs to be rearranged . In terms of natural light , the northeast side of is best suited. Then, in the morning hours, the plant will be illuminated by direct sunlight. They are not dangerous orchids, because they quickly hold on for a short time, until noon. When the heat hits in the afternoon, a shadow forms on the northeastern side that protects the orchid from the scorching sun.
In the case where the plants are located on the southern windowsill, it must be protected from overheating; it is protected with a light diffusing screen . It can be made of mesh fabric or mirror film.
From the scorching sun and rapid evaporation of moisture, phalaenopsis can be protected with louvers.
Phalaenopsis are not grown from the northern side. From a lack of light, it will not bloom.
See also article ⇒ How to plant orchid seeds at home.
Adjust temperature setting
Optimal heat for an orchid is within this range +15 0 – + 25 0 .
- In summer Phalaenopsis protects against heatstroke and excessive evaporation of moisture. If necessary, builds shading to protect the plant from overheating.
- In winter , it is necessary to ensure the temperature is not lower than +15 0 , otherwise frostbite will occur, after which the leaves die completely.
At temperatures below +150, phalaenopsis leaves wither from the cold.
Protect against drafts
If the leaves have withered due to strong gusts of wind, it is enough to rearrange the orchid in a cozy, protected place. At the same time, it is important to take into account that there should not be air stagnation there. In such an environment, pathogenic microorganisms develop well. It may happen that by saving the plant from drafts, conditions for the risk of diseases are created for it.
K. L. Kolosova – st.n. with. department
South American Plants
Botanical Garden, St. Petersburg.
Revise the humidification system
Most often, leaves wither from insufficient watering a. The difficulty is that it is impossible to establish a strict irrigation schedule. Watering frequency is selected individually. The interval between sessions is determined by the state of the substrate and the color of the roots.
The fact that it is time to water the phalaenopsis is evidenced by a silvery-white bloom on the roots.They are visible through the transparent walls of the pot.
- Another criterion for the next watering is the state of the substrate . It should dry out completely, but not turn into a crust. Drought can be the most likely cause of sluggish leaves. If you organize watering without drops of stagnant moisture and drying out of the substrate, then the problem of wilting will be resolved by itself.
Important! In addition to root irrigation, additional moisture evaporation must be created.For this, a wide and shallow dish with water is placed next to the orchid. As the level decreases, water is added. This technique helps to maintain the firmness of the leaves and reduce the frequency of root watering.
Available means for water retention in orchid tissues
If leaf wilting has just begun, and the reason is excessive evaporation, you can use glucose or sugar in order to retain moisture in the plant cells. For this, a solution is prepared:
- 1 teaspoon sugar in 200 ml (1 glass) water, or
- 10 ml of glucose per 5 liters of water.
A cotton pad (or a piece of soft cloth) is moistened in the solution and the leaves are wiped on both sides. After drying, a film forms which reduces evaporation. In addition, the nutrients serve as foliar feeding.
If these measures did not help, then you need to move on to more radical actions.
Phalaenopsis transplant. In what cases is it necessary. How to implement
Root problems
Sometimes the leaves of phalaenopsis die from the fact that the root system suffers.This can be due to excessive compaction of the substrate. Orchid roots receive nutrients differently from other plants. Substances are absorbed by the roots in a gaseous state from the air. From its lack in dense soil, the root system suffocates and cannot normally supply the ground part. To solve the problem, it is enough to transplant the plant into a new substrate with optimal aeration. .
In case of frostbite in winter or dry leaves in summer , the ground part of the orchid can be restored by replanting.But, first, the plant must be properly reanimated.
To do this, you need to perform the following work:
| Step 1 | Remove the plant from the pot and carefully examine the roots. Chances are good that the leaves have withered due to the poor condition of the root system. |
| Step 2 | Clean from substrate, if necessary wash in warm water. |
| Step 3 | Remove all damaged roots and leaves. |
| Step 4 | Sprinkle the sections with crushed activated charcoal or charcoal for disinfection. |
| Step 5 | If the orchid is left without leaves, the rest of the plant must be planted in fresh substrate and placed in a homemade greenhouse. It is constructed from a transparent plastic container or plastic bag. Every day the greenhouse must be opened for ventilation. When new leaves grow back, remove the protection.In the future, the rescued phalaenopsis is grown in the usual way. |
How to determine if a transplant is needed
If the withered leaves of the orchid did not recover after changing its location and watering, then the reason is different.
You can do this test:
- Move the rosette of leaves from side to side . It’s good if it moves. But when it sways freely and does not stick to the base, it means that the ground part has died.This most often occurs from waterlogging. Water accumulates in the axils of the leaves, and they rot away.
To save Phalaenopsis, he must be reanimated. First of all, the root system is restored with the help of growth stimulants:
- Heteroauxin,
- Kornevin,
- Succinic acid.
The roots are soaked in a solution of these preparations for several hours before planting.
Tip # 1 .Note! If the orchid has lost all its leaves, they can be re-grown if the roots are healthy.
Sometimes the opposite happens – the roots die, but the leaf rosette is still alive . In this case, all dead tissue is removed. The rest is soaked in a growth stimulant solution. In addition to the above-mentioned rooters, you can use “Epin – Extra”. It should be noted that the prepared solution must be used immediately, it quickly loses its beneficial properties in air.
Epin – Extra – stimulant for the restoration of leaves and roots.
Recommend this proportion:
- 1 ml of liquid preparation per 5 liters of water.
The water should be soft, free of alkaline salts, which reduce the effectiveness of the drug.
Read also the article ⇒ Growing an orchid of the Bulbophyllum variety: reviews, photos, reproduction.
Heading: “Question – answer”
Question No. 1 . Why did the phalaenopsis leaves wilted after I moved it to another place?
Answer.
Orchid is a very delicate plant, it reacts sensitively to any, even minor, interference. The wilting of the leaves is a “protest” against the undesirable changes for her. Most likely, after adaptation, turgor will recover.
Review
Anna Andreevna (Zagorsk)
I gave my orchid to a neighbor while on vacation. Upon my return, I received back limp and wrinkled leaves. I had to take care of it for almost 4 months.
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“Everyone makes his own step on the pedestal” – Sport – Kommersant
Olympic champion at 800 m about her performance at the first Games in her career.
– How are you feeling now?
– I’m so glad. Because she set a goal to win any medal, but she got the “gold”. Everything worked out today, everything worked out. And the mood, and the preparation, and the peak of the form, and the state of health, and the weather. However, I will not hide, I was afraid of force majeure circumstances at the distance. God forbid, I would have run into someone. But nothing happened. It all happened so quickly … I know one thing for sure: here the final race was easier for me than in Daegu.
– Did you realize in the morning that everything will be as planned in the evening?
– Everything was not clear in the morning.I remember that my legs were sluggish as never before, my general condition was not very good, my head ached and tears welling up. And I went to the start – and everything is fine. True, I tried my best to keep myself in good shape before the start. And the coach – Vladimir Kazarin – helped me a lot with this. To get into the vein, you need to be able to manage adrenaline. So it turns out that when my tears come before the start, it’s luck.
– When you ran behind the entire first lap, did you control all your opponents?
– Of course.I started running the way I need it. She did not rush and show frantic seconds at once. This tactic suits me better.
– Are you willingly or unwillingly using the tactics of Yuri Borzakovsky? He’s in the 2004 Olympic final in Athens, and you ran exactly the same here.
– I admire Yura, I am proud to meet him, but I have never consulted about my tactics. I just run the way it suits me. And the fact that we use the same technique with him is great.After all, not every runner or runner is capable of accelerating at the finish line as we can …
– Team tactics among Russian women, of course, were excluded?
– It doesn’t exist in our discipline. Here everyone is for himself, because he really wants to win. In addition, we train with different specialists, and they all have their own tasks. Everyone takes their step on the pedestal. In the finale, I started quickly. Probably, she was afraid that not only I would be able to switch to the finish line so quickly. But maybe someone else.All right.
– Who was feared more?
– Yes, I was afraid of everyone. Because the finals brought together the strongest 800m runners of the past few years.
– Is it easier or harder to become an Olympic champion than you thought?
– This is very difficult. You have no idea how much work, emotions was spent. And you can’t count the injuries. Therefore, I am grateful to all the specialists who helped me to approach the Games in excellent shape.
– What’s your next global goal?
– Rest globally.And now I will not think about the world championship in Moscow. Although, of course, I would like to speak at it. And no less successful.
Valeriya Mironova
| But, mainly, I’m tired of life in the city. Its monotony, my endless sluggish attempts to work and study. | ‘But mainly I was bored of life in the city, with its repetitions, my half-finished, half-hearted attempts at jobs and various studies.’ |
| Other results | |
| I was dormant for a while. Was pretty sluggish. | I have been in a state of hibernation for some time … a little inactive. |
| I woke up when we arrived, but initially lethargic and puzzled. | He was awake when we arrived, but initially lethargic and confused. |
| After trying different subjects of conversation, she directed him to the painting, about which he spoke very well, and listened to him attentively. | After trying various subjects of conversation, she got him upon painting, of which he talked very well, and she listened to him attentively. |
| Any ship attempting to land will be destroyed. | Any ship attempting to land will be destroyed. |
| Journalist Clayton M. McCleskey writes: “Mr. Koehler was isolated – a lonely leader trying to make Germany understand where it ranks in the world.” | Journalist Clayton M. McCleskey complained: “Mr. Koehler found himself isolated, a lonely leader attempting to push Germany to recognize the reality of its place in the world. ” |
| This was a man trying to get to the North Pole in a sleigh. | This was a man who attempted to go to the North Pole on a sledge. |
| With exceptional patience, the Special Rapporteur revised his draft articles on prevention in an attempt to eliminate all references to harm. | With extreme patience the Special Rapporteur had reformulated the draft articles on prevention, trying to delete all references to harm. |
| In May 1996, the NLD took another disastrous step by attempting to organize public rallies between 26 and 29 May. | In May 1996, NLD took another sinister step by attempting to hold a public rally from 26 to 29 May. |
| In short, we prefer to prejudge the outcome without even trying to work on it. | In short, we have preferred to prejudge the outcome without even beginning to work towards it. |
| Russia with its aggression opposed this desire, trying to test its strength. | That commitment has been challenged by Russian aggression. |
| Matt O’Brien posted a very nice post on the Atlantic, trying to explain why some countries performed well and others were downright weak.And I urge everyone and everyone to find out on their own. | Matt O’Brien at the Atlantic had a very good post that tried to explain why certain countries performed well or poorly and I encourage everyone to check it out. |
| According to one of the firms, the investment conditions in Iran are so incomprehensible, and this firm left without even trying to get compensation for its 60% stake in the company it had previously created. | According to one of the firms, investment conditions are unclear in Iran, and it left without even seeking compensation for its 60% share in a company it had created. |
| Not surprisingly, stock investors reacted to the outbreak of animal spirits by trying to get ahead of the likely growth in economic performance. | It is not surprising that equity investors have responded to the surge in animal spirits by attempting to run ahead of a possible uptick in economic performance. |
| I hope your spy informed you that four of my men died trying to arrest him. | I would hope that your spy also informed you that four of my six men attempted to arrest him and died in the effort. |
| Margaret gasped as she tried to speak, and when she spoke, her voice sounded muffled. | Margaret choked in trying to speak, and when she did it was very low. |
| But the pain at the base of the skull, which the girl felt when she tried to raise her head, indicated that this was not a dream at all. | But when she tried to lift her head, the pain at the base of her skull informed her of her wakefulness. |
| You have shown wisdom in trying to awaken these primal instincts. | You showed your wisdom in trying to reach that deep, underground, savage instinct. |
| Trying to assess the situation, he carefully examined the cotton shirt, into which the doctors had stuck his body, and looked for his clothes. | He took stock, examining the paper robe they had put him in. |
| Mrs. Boynton brandished her stick, tried to get up again, then collapsed into a chair. | Mrs. Boynton shook her stick, tried to rise, then fell back into her chair. |
| Dr. Harper, ”she said, trying to firm her voice. | Dr. Harper, she said, infusing her voice with intensity. |
| You will now lap the circuit, attempting to beat the time set by a Ferrari 458. | |
| Thought that I should retreat without even trying … | The thought that I must stand aside, not even try… |
| If the banks do not help us and Arnil and his entire company wants to evade, we will go bankrupt, that’s all. But without trying to save my own skin, I won’t give up, damn it! | If the banks won’t help us and Arneel and that crowd want to get from under, we’ll fail, that’s all; but not before I’ve had one more try, by Jericho! |
| Which means that maybe Franci … … didn’t do so badly trying to change herself. | It means that maybe Frenchy wasn’t so wrong to try and make something out of her self. |
| Having lived a life of manic depression and tried several times to commit suicide, Plath committed suicide on February 11, 1963 in her London apartment. | After living a life of manic depression and attempting suicide multiple times, Plath committed suicide on the 11th of February 1963 in her London flat. |
| Maimonides taught that the sages said that inspiration does not come to the Prophet when he is sad or languid. | Maimonides taught that the Sages said that inspiration does not come to a prophet when the prophet is sad or languid. |
| Trying to become a theater manager, he leased the Haymarket Theater in 1746. | Attempting life as a theater manager, he secured a lease on the Haymarket Theater in 1746. |
| However, upon entering the building and trying to find the Prime Minister, they were fired upon by four policemen. | Upon entering the compound and attempting to find the Prime Minister, however, they were fired upon by four policemen. |
| Although observations of live goblin sharks are scarce, their anatomy suggests that their lifestyle is inactive and sluggish. | Although observations of living goblin sharks are scant, its anatomy suggests its lifestyle is inactive and sluggish. |
| And the thought of coming to terms with death, letting the black panther gnaw itself on – was stupid, lethargic, unworthy. | To accept death, to let the black panther tear him to pieces, was stupid, weak, unworthy. |
| He was allegedly upset that the regiment retreated so quickly without trying to further delay the North Koreans. | He allegedly was upset that the regiment retreated so quickly without attempting to further delay the North Koreans. |
| After trying to leave, she discovers that someone has broken the wheel of her bicycle. | Upon trying to leave, she finds that someone has destroyed the wheel of her bicycle. |
| The nursing home sued her son directly, without even trying to collect money from Medicaid. | The nursing home sued her son directly, before even trying to collect from Medicaid. |
| Mohsen was arrested after breaking his bail agreement by attempting to flee the country. | Mohsen was arrested after violating his bail agreement by attempting to flee the country. |
| However, the Crusaders fought back by trying to place Laulala upright next to his right hand. | The Crusaders struck back though, with a try to Casey Laulala near the right hand upright. |
| Its façade consists of four Leo mythical figures holding columns, and two pilasters. | Its facade consists of four leonine mythical figures vyala, holding pillars, and two pilasters. |
| The façade has four pillars and two pilasters on one side of the main, oblong communal hall. | The facade has four vyala pillars and two pilasters on one side of the main, oblong community hall. |
| Above this is sitting a lion-in front of him. | Above this are seated lion-faced vyalas. |
| Yes, I am sluggish, and my anger is sluggish… Again I will see some boy, I will regret it, again I will be executed. No, it’s better this way … | And I am flabby, and my malice is flabby … I’ll see some little boy again, will have pity on him, will be punishing myself again … No, it’s better … better so! … |
| She, sluggish from lying down for a long time, was sitting at the table and writing something on a piece of paper. | Languid from lying down so long, she was sitting at the table, writing something on a scrap of paper. |
| Billy had not yet fully woken up and hardly perceived her reproaches, and the girl, warm and sluggish after sleep, was busy, looking under the mattress for her stockings. | Billy wasn’t awake enough to respond much to her shaming, and the girl was fussing around looking under the mattress for her nylons, moving slow and warm-looking after sleep. |
| Well, that’s a sluggish reaction. | Well, that’s a flaccid response |
| A sluggish, dull life devoid of any mental interest stretched out. | They now lived a dull, drowsy life, devoid of all intellectual interest. |
| Suddenly, without expecting it, Adam threw his hand up at random, and without aiming at anything, a flabby hand described a harmless arc in the void. | Suddenly to his own surprise Adam launched a wild, overhand, harmless swing which had neither force nor direction. |
| His own, exhausted by X-rays, sluggish, sick blood was still dearer to him than a fresh supplement. | His own sluggish, diseased blood, ruined by the X-rays, was still more precious to him than any fresh addition. |
| For two minutes he stood at the table in the same position, apparently deep in thought; but soon a languid, cold smile squeezed out on his lips. | He remained standing for two minutes in the same position by the table, apparently plunged in thought, but soon a cold and listless smile came on to his lips. |
| An inner tremor shook Lillian, but a sluggish, weak-willed nature forced her to restrain herself and be silent – why waste his time? | She paused half trembling and yet decided, because of her peculiarly nonresisting nature-why trespass on his time? |
| But Rench’s wife, a languid, apathetic woman, only wrapped herself in a large kerchief like a mummy, and he apparently set up his house incorrectly from the very beginning. | But Wrench had a wretched lymphatic wife who made a mummy of herself indoors in a large shawl; and he must have altogether begun with an ill-chosen domestic apparatus. |
| Minnie was filling a mug from a bottle, and as it filled, a sluggish foam rose. | Minnie was filling the tankard from the bottle, a dull frosting pacing the rise of liquor within the metal. |
| When the cell membrane has a low turgor pressure, it is flaccid. | When the cell membrane has low turgor pressure, it is flaccid. |
| The lower this percentage, the lower the rate of maturation and the more sluggish the disease. | The lower the percentage, the lower the speed of maturity, and the more indolent the disease. |
| It would be great if it was dried, as the cod is dried. | It’d be nice if it was dried, because codfish is a dried fish. |
| If it was dried fish or bananas… | If it was dried fish or bananas … |
| No, the pork was dried with salt. | No, these pigs were cured with salt. |
| Thus, Iberico jamon is a dry cured ham obtained from cattle of these breeds. | As such, jamón ibérico is the dry-cured ham produced from livestock of said breeds. |
| Depending on the recipes, it can be with pickled, steamed, toasted or dried foods. | According to the recipe difference, seasoned, parboiled, battered and fried are the typical brochettes. |
| Lokhiro looked away from the food, which consisted of flatbread and jerky, which he greedily devoured. | Lohiro glanced up from the meal of pan bread and dried meat he had been devouring. |
| The aroma of stew of turnip stew, onion and jerky drifted over the camp. | The smell of a stew made from turnips, onions, and dried meat settled over the camp. |
| Then he made a fire, cooked a stew of dried meat and went to sleep for a couple of hours. | He prepared a small fire and made some broth from his dried meat before snatching two hours’ sleep. |
Zakharova called Kosovo the “black hole” of Europe
https://ria.ru/20210628/zakharova-1738880529.html
Zakharova called Kosovo the “black hole” of Europe
Zakharova called Kosovo the “black hole” of Europe – RIA Novosti, 28.06.2021
Zakharova called Kosovo the “black hole” of Europe
Foreign Ministry spokeswoman Maria Zakharova called Kosovo a failed European project. RIA Novosti, 28.06.2021
2021-06-28T12: 11
2021-06-28T12: 11
2021-06-28T14: 38
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MOSCOW, June 28 – RIA Novosti. Foreign Ministry spokeswoman Maria Zakharova called Kosovo a failed European project. According to her, the new Kosovar authorities “preach open political extremism.” In March, deputies of the self-proclaimed republic’s legislature elected Albin Kurti, leader of the radical Self-Determination movement, as prime minister. It advocates limiting Belgrade’s ability to influence the situation in the region.Zakharova believes that “the western project” Kosovo “failed,” despite the funds and information and political support “pumped into” it. The Foreign Ministry spokeswoman also recalled that in 2013 the so-called Brussels agreements were agreed. Serbia, the diplomat claims, has complied with its part of those decisions, while the other side refused to provide a key condition – the creation of the Community of Serbian Municipalities of Kosovo, endowed with broad administrative and executive powers.”Moreover, the current Kosovo authorities do not want to discuss this issue at all. Brussels cannot do anything about it, limiting itself to sluggish calls without concrete steps to force the Kosovo Albanians to abide by their obligations,” Zakharova added. The Kosovo Liberation Army and the Serbian army and police led to the bombing of Yugoslavia (which at the time was Serbia and Montenegro) by NATO forces. On February 17, 2008, the Kosovo-Albanian structures in Pristina unilaterally declared independence from Serbia.The decision is not recognized by Serbia, Russia, China, Iran, Spain, Greece and a number of other states. Belgrade in 2011 began negotiations on the normalization of relations with the Kosovar Albanians, mediated by the EU.
https://ria.ru/20210322/kosovo-1602402435.html
https://ria.ru/20210517/kosovo-1732365760.html
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12:11 28.06.2021 (updated: 14:38 28.06.2021)
Zakharova called Kosovo the “black hole” of Europe
90,000 First steps in wine tasting
Professional tasters always follow a strict sequence. First, the appearance of the wine is described, then its aromas, and only after that – the taste characteristics.
Your first tastings are best done in the company of someone with such experience.However, you can cope on your own, armed, for example, with our advice. First of all, several points should be noted that are very important in order for the tasting to be successful.
Furnishing . Wine can be perceived in different ways, depending on certain factors. In addition to lighting and temperature, even the time of day matters, your well-being and mood.
The air during tasting should be fresh, free of foreign odors, lighting – daytime or close to it.The temperature of both the wine itself and the room is very important. If it is too cool, the aroma of the wine will not develop well, if it is too hot, alcohol vapors will distort the bouquet.
Glass . A clean wine glass is required for tasting. It should be in the shape of a tulip: narrower at the top, wider at the base. This will allow you to capture the aroma as much as possible and prevent the wine from spilling out when you rotate it. For tasting white wines, glasses of a smaller volume are usually used, for red – larger ones.The way you hold the glass is of particular importance when tasting. Never grab the top of it – just the leg. Otherwise, the warmth of your hand will further heat the wine.
To start . Read again the description on the bottle label, so that then, during the tasting process, try to find the peculiarities of the variety declared by the manufacturer in the wine. Pour some wine into the glass. If you are tasting several wines, start with a light white, then move on to heavier red wines.This will help to keep your taste buds sensitive, making it possible to more accurately evaluate each wine. A sip of water between tastings will also help preserve your taste.
Visual Analysis
Take a close look at the wine. First, examine its surface, how shiny it is. Then tilt the glass a little, look through it at the light or at a sheet of white paper. What do you see? Is it clear or cloudy? How intense is its color? Wine color should be discussed separately.On this basis, all wines are divided into three main categories: white, rosé and red. However, the color spectrum of each of them changes with age.
Red wines . They vary greatly in color. Young wine is usually bright raspberry with reddish brown tints around the edges. Older red wine can range from reddish brown to brick in color.
White wines . The shades of these wines range from pale straw and lemon to deep golden amber tones.
Smell analysis
Determination of aromas is the most difficult part of tasting, it requires a lot of concentration. It should also be reminded that you cannot taste wine with a stuffy nose.
Firmly holding the stem of the glass, gently but vigorously rotate the wine for 10-20 seconds. This will allow him to ventilate and get oxygenated. Getting on the walls of the glass, it will reveal its bouquet.
Wine contains more than 500 aromatic components, there are many types of its aromas.For example, here are the main ones:
- fruit and berry aromas – black currant, apricot, peach, apple, raspberry, cherry, cherry, strawberry, lemon, fig, prune, etc .;
- flower aromas – rose, wild rose, violet, jasmine, acacia, iris, peony, etc .;
- vegetable aromas – grass, mint, eucalyptus, moss, mushrooms, etc .;
- spicy aromas – cloves, pepper, ginger, etc.;
- empiromatic aromas – toasted bread, tobacco, roasted coffee, roasted almonds, etc.;
- balsamic aromas – resin, incense, pine, etc.;
- animal aromas – meat, game, fur, musk, etc.
So, after making a few rotational movements, tilt the glass and, if possible, lower your nose into it. Take a deep breath as if you were sniffing a flower. The aromas can be completely different depending on how much your olfactory organ will be dipped into the glass. However, there is no special inhalation technique. Some wine connoisseurs prefer to sniff by inhaling quickly two or three times.Others take one deep breath or one nostril.
Try to interpret the sensations that you can catch. Also, pay attention to whether the wine has a light or rich aroma, simple or polysyllabic. After a few breaths, your nose will get used to it and begin to feel less nuanced. In this case, it is better to pause, set the glass aside, and continue after a few minutes. You will probably notice that in the upper part of the glass, the aromas are more floral and fruity, deeper, closer to the bottom – they become richer and richer.Try to discover the full spectrum of these scents.
Take a sip and taste
This is the last step and is only done after you have examined the wine and smelled its aromas.
Take a sip of the wine, but do not swallow it. Let the guilt spread over your tongue from front to back, side to side. Try to chew on it. If you succeed, blow some air into your mouth through your covered lips.Its ingestion will help release the taste of the wine and its aromatic components.
In the mouth, the wine heats up, begins to exude new aromas, which are perceived by the nasopharynx. The nose senses the subtlest odors, the mouth makes it possible to feel the components of the taste.
The tip of the tongue determines the sweetness, the inner side of the tongue and its upper edges feel acidity. The back of the tongue reveals bitterness and alcohol, the inner side of the cheeks – tannins. At this moment, all the virtues of wine are finally manifested.You will be able to analyze the full range of taste sensations and consistency. A good wine will always be harmonious and balanced in terms of sugar, acidity and astringency. After it, a complex and long aftertaste will definitely remain.
At this moment you can, as it were, touch the wine. This, of course, does not mean that you should dip your finger in the glass! When tasting a wine, the touch with the wine happens in your language. How do you feel about it? Is it soft or aggressive? Does it have refreshing energy? Or is this wine flat and sluggish? The tannins found in red wines can give the wine a viscous or silky feel, depending on how ripe they are.
At this point, you can spit it out (especially if you are tasting several wines) or just drink the wine, but be sure to smell and remember the aftertaste.
Overall score
After the aftertaste has passed, ask yourself what is your overall impression of the wine. Did you like it? Are all components balanced? Was this wine subtle or rough, simple or complex? You can write down your feelings.
We have told you about the basic principles of tasting.It should be noted that the assessment of the taste of wine is always very individual. It cannot be described unequivocally correctly or incorrectly. Each person’s taste is unique and different from the feelings of other people.
Yes, and the person himself, throughout his life, trying new wines, changes his initial preferences, his taste preferences change. His sense of smell and taste also become different. So, in the assessment of guilt there can be no absolute truth or absolute error.
Where to start?
You can carry out the first practical tasting experiments as follows.Buy several bottles of good wine from a specific category. For example, if you prefer red wine, these could be: Merlot, Cabernet Sauvignon, Pinot Noir, Carmenere and Syrah (Shiraz). It is better that these are young wines, as the taste and aroma of the wines change during the aging process.
Your task will be to find the differences in taste between these wines, to discover the color nuances inherent in each sample, to grasp the aromatic palette. Pouring wine into a glass, read again its description and try to find those features that are characteristic of it.
For example, the basic tones of Cabernet Sauvignon are black currant, blueberry, ginger and green pepper.
Merlot has fruity and berry aromas of cherry, raspberry, blackberry and plum.
In Pinot Noir you will smell flowers and herbs: violets, jasmine, lavender, eucalyptus and mint, as well as tones of red berries.
In Carmenere try to find notes of black currant, figs, prunes and green pepper.
