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Nobody wants to die, but it does happen. Do not neglect to plan something for your family because you do not want to think about this eventuality. The information in this article offers insight and advice for making the most of life insurance.
Look at your family's needs and lifestyle when determining the life insurance you need. Choose a policy adapted to the specific needs of your family after you are gone. When something like this does happen, it's good to know that your policy will be there to provide for their loved ones and cover the monetary damage.
Make sure you have adequate coverage when buying life insurance, enough to take care of your loved ones. You policy should cover all your debts and expenses for your spouse and your children.
Before you buy life insurance, compare prices with other companies. The price differences between companies can vary widely, as much as 50%. Check the coverage prices online at comparison websites as well as the insurance companies' official sites. Also, choose a policy whose price takes into account your medical history.
Bungee jumping, skydiving and scuba diving in your free time, can give your life insurance provider cause to increase your premiums. Additionally, some occupations, such as helicopter pilot or a race car driver, can be considered high risk by your insurance provider, and they will increase your premiums accordingly.
Try getting your life insurance from a financial adviser and not an insurance broker. Insurance brokers earn commissions from the sales of life insurance policies. Financial advisers earn the same fee, no matter which policy you decide on. This is why financial advisers often do not push you to buy and they will be more honest.
If your job is high risk, or if you have any hobbies that would be thought of as high risk, you need to disclose them to the life insurance company. It is going to cost you more money, but it is going to keep you from becoming ineligible for coverage if the insurance company finds out on its own. It might be considered fraud if you withhold the information.
Try improving your health to get cheaper life insurance. Since healthier people have a longer life expectancy, they get better deals.
Watch out for any signs of shadiness from the person you are working with. If an agent makes outlandish claims, or ignores the rating agencies, it's best to work with someone else, and to mention their claims to their supervisor.
When buying the life insurance policy, an independent broker will often be able to offer a better deal than a firm that solely provides insurance. Independent brokers genuinely offer you products wide in range, as opposed to a products that are specific to a firm. Because you are obligated to your life insurance policy, it's important that you look around for the best one.
As was said before, no one wants or plans to die, but it will happen. It happens a lot sooner than people might plan on. Disastrous events for your family could occur by not planning properly. The following article will enable you to provide for your family, even in the event of an untimely death.


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Consider starting your planning sessions prayerfully. Post this prayer with your VBS planning materials to remind yourself that VBS belongs to God, and watch Him work through you, the volunteers, and children. Seek His guidance and praise Him often.
Lord God, we ask Your blessing on Barnyard Roundup VBS. Kids and families need to hear the message that "Jesus gathers us together." The Lord is MY shepherd! Your care, provision, leadership, forgiveness, and salvation in Psalm 23 are the "now and forever" good news we all need! Provide Your presence and strength for all of us who are planning VBS 2016. Send Your holy angels to guard and protect us that Satan does not hinder the journey of Your Word as proclaimed in Barnyard Roundup to the hearts and lives of children, volunteers, and families. Lord, in Your mercy, hear our prayer for Jesus' sake. Amen.

subset: commoncrawl


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Mould and mycotoxins risks in feeds
Drought and low growth seasons tend to see a range of less common feed ingredients being used. These include older hay, alternative co-product feeds, and a high volume of failed cereal and canola crops. It is important to understand potential risks and the upsides of using these types of feed.
Oxygen is the enemy in silage production
Poorly stored hay, silage and/or other high moisture feeds are at risk of spoilage by microbes (e.g. yeasts, bacteria and moulds). Silage made from mature crops has a high risk of spoilage as it is hard to keep oxygen out.
Alternative crop hays are often at risk of becoming mouldy in storage because of their high moisture content and higher microbial load than normal hay sources.
White moulds can be dangerous. Consider testing your forage to reduce the impact of toxins on your livestock. If feed is at risk of fungal impact, mycotoxins management tools could be used to alleviate the symptoms.
- Intake of sugarcane silage by lactating cows is less than other forages
- Low feed intake, which can occur with simple fungal impact, or with toxin load
- Rough hair on the coat
- Slightly arched back
- Swollen hocks and laminitis
- Digestive upsets such as diarrhoea and/or rumen stasis
- Mucous in manure
- High somatic cell count
- Excess salivation
- High rate of abortion or foetal resorption
- Broader infertility issues.


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So wow, I can't believe this is link up number 20! A massive thank you for all of you for sticking with us every week and for those newbies always joining! You are very welcome! I love watching all your videos every week and learning all about everyone. I never fail to get inspiration or learn something new out of every video I watch.
If you are new to this linky, it goes live every Sunday morning and you can link up any video you like, old or new. You can link directly to the video or the blog post, it's up to you. If you have embedded your video into a blog post feel free to add the badge to raise awareness of the linky.
I promise to watch and share on social media every one of your videos. It would be great if you could watch and comment and thumbs up a few others in the link up – that's how we share the love! Remember that commenting on other videos is not only a nice thing to do for a fellow vlogger, but it's also a great way of advertising your channel, as other commenters will see your name. And no one likes a link dumper 😉
Last week I highlighted Becky-Goddard's Hill video as she has a new linky and I was so pleased to see that some of you linked up. I hope that I still have a few days left to add mine. So this week's favourite video comes from Maria Loves with her video for the link up – her top ten colours! We have similar tastes I think so she chose colours I would choose myself, with one very unsual 'colour' – take a watch to find out.
[inlinkz_linkup id=693630 mode=1]
If you have embedded your video in a blog post feel free to add the badge so we can raise awareness of the linky. Sorry last week the code was wrong, I noticed when I saw it on someone's blog! It's fixed now!


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Today, the Founder of Peace and National Unity - Leader of the Nation, President of Tajikistan Emomali Rahmon accompanied by the Mayor of Dushanbe Rustami Emomali inspected several locations in the city. The aim of the tour was to check on the progress made in implementing the presidential instructions, including the construction of the Central Mosque, landscaping, planting of seedlings, environmental protection and preparations for the international Navruz celebrations.
Today, in the early days of the beautiful spring season and in order to worthily celebrate International Navruz holiday and expand landscaping, President of the country Emomali Rahmon and Dushanbe Mayor Rustami Emomali attended a tree-planting campaign in the territory of the Central Cathedral Mosque of Dushanbe.
Firstly, the head of state Emomali Rahmon inspected the state of progress in construction of Central Mosque and construction site of the Islamic Institute of Tajikistan named after Imom Azam - Abu Hanifa Nu'mon Ibn Sobit and planted several maple seedlings on the territory of the mosque.
While planting the seedlings on the territory of the Central Cathedral Mosque, the inspections moved on to the Navruzgoh Park. The park encompassed an area of 146,000 square meters and includes multiple varieties of trees and other greenery.


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Fill the form (or) Call +91-9311222578
Corporate packers movers delhi services offered by many movers and packers is of recent origin. Today since the business are growing day by day, we see more and more companies shifting and expanding their base and so such services have come into play and are more and more in demand. The services provided are so good that the company looking for relocation forgets all its tensions and worries. They simply contact the office shifting company delhi either personally or through phones or internet and half the battle is won.
The experts from the engaged company will visit the premises and will get to know about your needs and requirements and then will provide you with a plan and quotation. You will be surprised to see that the plan is not only flexible and really impactful also the price quoted is much less than you expected. The corporate relocation services provided by such companies also offer the company to operate at odd hours i.e. when there is a public holiday or there is a weekend. Also there would be no need to spare manpower from the company's. The staff employed is well trained and will single handedly carry out the entire move procedure with great efficiency.
Also the service providing company is going to bring its own packaging material and will also arrange for the transport. Many companies have got their own fleet of vehicles while some have contact with various transport companies. They will also provide you with services like car and other vehicle transportation. And will also provide you with insurance and warehousing facility if there emerges any need. One should not feel that such services are meant only for the well off sections of the society rather these services are meant for the benefit of the mankind as a whole and so are designed in a way to suit everyone's need and pocket.


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Store hours may vary.
Please contact Lasalle Court Mall directly for shopping hours information.
Lasalle Court Mall offer more than 14 brand name stores. Lasalle Court Mall is located in Sudbury, Ontario - 900 Lasalle Blvd, Sudbury, ON P3A 5W8 (GPS: 46.522407, -80.96125).
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Lasalle Court Mall - Sudbury Area, Ontario P3A 5W8. Click on the button and get directions to Lasalle Court Mall. Mall/Shopping center is also displayed on the map with gps for your navigation.
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Tisserand 3 Step Ritual To Destress Kit Essential Oil Blend Boxed Gift Set
A 3 step remedy to de-stress your mind and body. Infused with a soothing blend of geranium, orange and nutmeg, it's perfect for helping to rebalance and calm.
Featuring a body oil to soften and calm the skin, a mist to lift the spirits and melt away stress, and a roller ball to ease tension, it's ideal for anyone suffering with stress or anxiety.
- Total De-Stress Massage & Body Oil 9ml
- Total De-Stress MoodFix Mist 9ml
- Total De-Stress Pulse Point Roller Ball 10ml
Part of our Award-Winning Wellbeing collection, the 3-Step Ritual To De-Stress box is the perfect go-to companion for busy lives. Containing Massage & Body Oil for the body & mind, Moodfix Mist for your room and Pulse Point Roller Ball for instant comfort.
Expertly blended with 100% natural pure essential oils to help reclaim your calm. Let the tranquil fusion of Geranium, Orange and Nutmeg help you slow down and breathe when life is getting on top of you. Geranium helps level the senses, while uplifting Orange lightens the weight of a heavy body and mind. Nutmeg creates a supportive feeling of comfort, helping you to let go of stress and enjoy a sense of calm.
Sparkle and Shimmer,
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Please note: Unless specifically stated otherwise, it should be assumed that no product sold on this website is suitable for use by persons under 18 years of age. All prices are in £Sterling (UK Pounds). Waste Electrical & Electronic Equipment Regs 2006: For recycling info please click here.
Copyright © 2008-2019 Sparkle and Shimmer


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There's something about playing guitars and riding motorcycles that seems to go hand in hand. Over the years there have been several incredible musicians who've enjoyed the freedom of riding a motorcycle on the open road.
Artists including Bob Dylan, Neil Young, Alice Cooper, Gene Simmons, Willie Nelson, Duane Allman, Jimi Hendrix, and Kid Rock have all had their share of wind therapy. Many of these musicians rode motorcycles in their free time, but also had major influences on their music. Many musicians wrote songs about motorcycle rides ('Born to be Wild' – Steppenwolf), included motorcycles in their album covers, music videos, and even their on stage performances (Judas Priest). Check out this short, but sweet list of influential musicians that also rode motorcycles. Let us know in the comments your favorite musicians who ride and if any of these artists on the list surprised you. Check out the photos below:


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There's two dimensions to life, one mental, one physical.
In the physical dimension all things are so interconnected as to be one whole thing. In the mental dimension the same thing applies. The observable universe is God, the mind that looks out from you onto the material world which is the body of God is also God.
In the beginning there was nothing, so God created the universe out of himself. Likewise there was no one else so God peopled the universe out of himself.
All is God.


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door and window stretch wrap machine
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job is a very fast moving and volume program which had been earlier managed by 6 to several employees. Now we are able to run it with less than two people," states Joe McGillivray, Powerful Group co-owner, who actually didn't believe his company could afford collaborative automation."When
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Inside the same period, the door and window stretch wrap robot selections both the hygienic wipes as well as the saline solution, then places them to the clamshell."I also
assumed that I would require an extremely costly engineer to aid me get these worked out. I am no expert and within an hour or so of shipping, I needed them set up ready to go basic applications," provides McGillivray. He quotes the return in the preliminary method to get lower than sixty days.
Some of the substantial time and resource cost savings adding to rapid Return on investment was the truth that the UR robots didn't will need basic safety protecting."While the robots'
safety features continue to keep our workers safeguarded, they're allowed to practically interact with the door and window stretch wrap robot. That's a huge cost savings for us promptly, being forced to manufacture, and costs to manufacture safety enclosures.
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Original Research ARTICLE
Photosynthetic acclimation responses of maize seedlings grown under artificial laboratory light gradients mimicking natural canopy conditions
- Institut für Allgemeine Botanik und Pflanzenphysiologie, Lehrstuhl für Pflanzenphysiologie, Friedrich-Schiller-Universität Jena, Jena, Germany
In this study we assessed the ability of the C4 plant maize to perform long-term photosynthetic acclimation in an artificial light quality system previously used for analyzing short-term and long-term acclimation responses (LTR) in C3 plants. We aimed to test if this light system could be used as a tool for analyzing redox-regulated acclimation processes in maize seedlings. Photosynthetic parameters obtained from maize samples harvested in the field were used as control. The results indicated that field grown maize performed a pronounced LTR with significant differences between the top and the bottom levels of the plant stand corresponding to the strong light gradients occurring in it. We compared these data to results obtained from maize seedlings grown under artificial light sources preferentially exciting either photosystem II or photosystem I. In C3 plants, this light system induces redox signals within the photosynthetic electron transport chain which trigger state transitions and differential phosphorylation of LHCII (light harvesting complexes of photosystem II). The LTR to these redox signals induces changes in the accumulation of plastid psaA transcripts, in chlorophyll (Chl) fluorescence values Fs/Fm, in Chl a/b ratios and in transient starch accumulation in C3 plants. Maize seedlings grown in this light system exhibited a pronounced ability to perform both short-term and long-term acclimation at the level of psaA transcripts, Chl fluorescence values Fs/Fm and Chl a/b ratios. Interestingly, maize seedlings did not exhibit redox-controlled variations of starch accumulation probably because of its specific differences in energy metabolism. In summary, the artificial laboratory light system was found to be well-suited to mimic field light conditions and provides a physiological tool for studying the molecular regulation of the LTR of maize in more detail.
Plants need to cope with natural variations in illumination and temperature during their vegetation period. The major process affected by both parameters is photosynthesis, which is both light- and temperature-sensitive due to the tight coupling between light and dark reaction (Hüner et al., 1998; Blankenship, 2002). Photosynthesis, therefore, represents an attractive target for biotechnological improvements in order to promote growth and yield of crop plants (Long et al., 2006; Zhu et al., 2010).
Natural illumination of plants is highly fluctuating in intensity and quality mainly because of seasonal and daily periodicity and of short-term disturbances caused from, e.g., clouding or leaf movement. Beside these abiotic influences plants themselves create strong light gradients especially within dense plant populations where leaves of neighboring plants shade each other and compete for light. This competition for light results in strong decreases in light intensity and a relative enrichment in far-red wavelengths within the canopy (Terashima et al., 2005). The latter effect is sensed by photoreceptors which control photomorphogenesis or shade avoidance responses (Ballare, 1999; Smith, 2000). The enrichment in far-red light wavelengths has an additional strong impact on photosynthesis since it can be partly used by photosystem I (PSI) but not or less effectively by photosystem II (PSII) causing imbalanced distribution of excitation energy between the photosystems. During evolution plants developed a number of responses which optimize light utilization under these variable conditions (Aro and Andersson, 2001; Blankenship, 2002). Developmental responses at the whole-plant or leaf level include variation of photosynthetic capacity by modification of leaf thickness (e.g., sun and shade leaves) which take place within weeks and months. They act in concert with more dynamic molecular acclimation mechanisms at the chloroplast level which work within the range of minutes to several hours. These physiological responses are fully reversible. In the short-term functionality of the photosystem antennae is adjusted by important regulatory processes such as non-photochemical quenching for pH dependent dissipation of excess excitation energy under high light as well as state transitions – a posttranslational modification, which counteracts imbalanced excitation of the photosystems under low light. In the long-term, changes in gene expression control an adjustment of photosystem stoichiometry which work in the same functional direction as state transitions, but create a longer lasting effect (Melis, 1991; Anderson et al., 1995; Allen and Pfannschmidt, 2000; Haldrup et al., 2001; Allen, 2003; Holt et al., 2004; Pons and de Jong-Van Berkel, 2004; Kanervo et al., 2005; Walters, 2005; Eberhard et al., 2008; Horton et al., 2008; Kargul and Barber, 2008). These different responses generate a hierarchical framework which countervails the high variability in illumination and optimize photosynthetic electron transport (Frenkel et al., 2007; Dietzel et al., 2008). Therefore, it was proposed that manipulation of photosynthetic acclimation might help to improve photosynthesis (Horton et al., 2001; Murchie et al., 2009). In general, photosynthetic acclimation appears to be a chloroplast-autonomous regulation being independent of photoreceptors (Walters et al., 1999; Fey et al., 2005a); however, because of the numerous interactions in the plant cellular signaling network indirect photoreceptor and hormone influences must be also taken into account (Boonman et al., 2007, 2009).
Photosystem stoichiometry adjustment could be found in shade- and sun- as well as in mono- and dicot plants (Chow et al., 1988, 1990a; Pfannschmidt et al., 1999a; Fan et al., 2007) and appears to be an ubiquitous acclimation response in photosynthetic organisms (Melis, 1991; Allen and Pfannschmidt, 2000). Recent studies with the model organism Arabidopsis thaliana have revealed that the thylakoid-associated kinase STN7 is a key regulator of this long-term response (LTR; Bonardi et al., 2005; Pesaresi et al., 2009). Using knock out mutants as negative control it could be demonstrated that state transitions and LTR provide a physiological advantage under permanently changing conditions resulting in enhanced growth and seed production (Bellafiore et al., 2005; Frenkel et al., 2007; Wagner et al., 2008). Furthermore, systems biology approaches revealed that photosynthetic acclimation responses not only reconfigure the photosynthetic apparatus to the instantaneous light environment but also trigger a metabolic reprogramming which coordinates the energetic demands with the light harvesting efficiencies of the plant (Bräutigam et al., 2009; Frenkel et al., 2009). The studies demonstrated that starch accumulation in Arabidopsis significantly differs between different light quality acclimation states and, thus, represents a well-suited reporter for the metabolic reprogramming that finally controls plant growth efficiency (Bräutigam et al., 2009).
Maize is one of the world's most important field crops and improvement of its photosynthetic yield would be of great interest (Zhu et al., 2010). Maize fields generate tall and dense plant stands during its vegetation period, which contains strong light gradients and, hence, a high degree of photosynthetic acclimation should be expected. Recent observations demonstrated that maize exhibit a high ability for photosynthetic acclimation when grown under different white light (WL) intensities in growth chambers. Variations in antenna structures and LHC protein accumulation were found both in mesophyll and bundle sheath chloroplasts (Drozak and Romanowska, 2006).
Here, we analyzed the degree of photosynthetic acclimation in maize grown under low-intensity artificial light qualities, which preferentially excite either PSI or PSII. This test system was successfully used previously for understanding redox-controlled photosynthetic acclimation in C3 plants like mustard, tobacco and Arabidopsis (Pfannschmidt et al., 1999a, b, 2001; Fey et al., 2005b; Wagner et al., 2008). Light quality acclimation to artificial light quality gradients could be demonstrated to be efficient and beneficial in various C3 species (Chow et al., 1990b; Hogewoning et al., 2012). In order to test the corresponding response of the C4 plant we determined chlorophyll (Chl) a fluorescence, Chl a/b ratios, transcript accumulation of plastid gene psaA (encoding the core protein of PSI), phosphorylation state of light harvesting complexes of PSII (LHCII) and the ability to perform state transitions. The detected acclimation ability was compared to that of samples harvested from maize plants grown under natural conditions. Our study indicates a high ability of maize to acclimate to light quality gradients in low-intensity light conditions in a lab based system providing a tool for analyzing the potential of the maize LTR for biotechnology applications.
Light Gradients and Photosynthetic Acclimation of Maize Under Field Conditions
Light intensity within plant populations decreases as a function of the leaf area index (LAI) while at the same time an enrichment of far-red wavelengths occurs (Smith, 2000; Hirose, 2005). We wanted to test if we could use a laboratory light set-up to study acclimation responses of maize seedlings to natural light gradients. In order to have a reference we analyzed the course of light gradients in a maize field trail by determination of light intensity and spectrum at different heights (Figure 1) of full-grown maize plants. 80% of the light was absorbed within the top 10–20% of the field, all lower parts obtained low light intensities with a strong enrichment in the far-red range (Table 1). Thus, even under sunny conditions the majority of the leaves in the field perceived only a sub-saturating photosynthetic active radiation (PAR) with a three- to four-times higher proportion of far-red light. To analyze effects on the photosynthetic performance of the plants we harvested plant leaf samples from the top, the middle and the bottom region of the stand. To include possible effects of the leaf blade position and the developmental gradient in chloroplast biogenesis we studied three segments of the leaves (basal, middle, and top segment; Figure 2A). In order to study the potential light acclimation we probed the Chl a/b ratio as indicator for changes in the amount of LHCII and the Chl fluorescence value Fs/Fm as physiological marker for the LTR, This value represents the fraction of light which is absorbed by the PSII antenna and subsequently lost by constitutive dissipation via fluorescence and heat emission in the steady state. Photosystem stoichiometry adjustment creates significant changes in this parameter (Pfannschmidt et al., 2001). The Chl a/b ratio was found to be significantly higher in level 1 than in level 2 and 3 (Figure 2B) while the measured Fs/Fm values were lower in level 1 and increased step-wise toward level 2 and 3 (Figure 2C) with a significant difference in section 2 of level 1 and 3. Effects of the leaf gradient were hardly detectable with the exception of segment 1 in level 2 which displayed a slightly more pronounced increase in the Fs/Fm values than segments 2 and 3. The starch content was found to be relatively stable at the different levels of sampling (Figure 2D) with a weak tendency to increase toward the bottom level which, however, was only significant in segment 2 of level 3. In summary, the measured parameters Fs/Fm and Chl a/b demonstrated typical changes between top and bottom level being indicative for a functional LTR in maize under field conditions.
FIGURE 1. Natural illumination conditions within the maize field used for sampling. Left picture displays the plant stand used for measurements. Light intensities determined at indicated heights within the stand are given in the middle panel. Corresponding light spectra (indicated by the same color) at the respective position are shown on the right. Open: intensity and spectrum of full sunlight outside the field. PAR: photosynthetic active radiation. FR: far-red region.
FIGURE 2. Analysis of field grown maize. (A) Left panel displays height of leaf harvesting. Magnification on right defines the analyzed sections of the respective leaves. Three to five leaves of each level were harvested from different individuals distributed randomly over the field. (B) Chl a/b ratio, (C) Chl fluorescence, and (D) starch contents in field grown maize. Story of harvest is given on the left, respective leaf segment analyzed on top. Acclimation parameters Chl a/b and Fs/Fm were determined as described in Section "Materials and Methods." Starch content is given relative to the fresh weight (fw). Data represent means from at least three independent biological replicates, standard deviation is given. A Student's t-test was performed to test significance of differences (*P < 0.05, **P < 0.01, ***P < 0.005).
Photosynthetic Acclimation of Maize Seedlings to Light Quality Gradients
Recently, we described the physiological and molecular acclimation of the dicotyledonous C3 plants A. thaliana, Nicotiana tabacum, and Sinapis alba grown in a light system which preferentially excites either PSI or PSII (Pfannschmidt et al., 2001; Fey et al., 2005b; Wagner et al., 2008; Steiner et al., 2009). Here, we used the same set-up to analyze photosynthetic acclimation in the monocotyledonous C4 plant Zea mays. Seedlings were grown under WL as control and under PSI- or PSII-light (PSI, PSII) in order to test their ability to perform a LTR. Additional controls were shifted between the PSI- and PSII-lights (PSI-II; PSII-I) to test the reversibility of the response, an important criterion to distinguish between acclimation and development. The general appearance of the plants grown under the different light regimes did not reveal major morphological differences which is consistent with earlier observations in C3 plants (Figure 3A). For detection of a LTR the second true leaves of 10-day-old plants were used for measuring the Fs/Fm parameter by video imaging (Figure 3A). In this experimental set-up PSI-light acclimated plants absorb more light than they can use and dissipate the excess as fluorescence which usually is significantly higher in the steady-state than that from PSII-light acclimated plants. A deficiency in performing a LTR, however, is indicated by a loss of significant changes in Fs/Fm as observed in the stn7 mutant of Arabidopsis (Bonardi et al., 2005). WL plants exhibited a low Fs/Fm value while PSI plants displayed a high one. After a shift from PSI-light to PSII-light the Fs/Fm value in the seedlings declined. In PSII-plants it was low and increased after a shift of the plants to PSI-light. This result is consistent with earlier measurements of Fs/Fm values in C3 plants indicating that maize seedlings are able to perform a LTR in the same manner.
FIGURE 3. Chl fluorescence of maize seedlings acclimated in long-term to different artificial light regimes affecting the redox state of photosynthetic electron transport. The growth light regime is indicated next to each leaf. (A) Left: appearance of leaves of differently acclimated plants under white light. Right: Fs/Fm values of the same leaves as detected by a CCD camera for Chl fluorescence detection given in false colors. Color bar on the right indicates the measured values. First and second true leaves were used for all analyses, the figures display representative samples. (B) Leaf sections used for more detailed analyses (compare Figures 4 and 5) were selected according to the field sampling.
2D images of the Chl fluorescence revealed some variations in the fluorescence values between different areas of the leaf blade (Figure 3A) which could be caused by the characteristic developmental gradient of the plastids in young monocotyledonous leaves which is more pronounced than in fully developed leaves as investigated in Figure 2. To examine this in more detail leaf blades were dissected into three segments containing young plastid from the leaf base (segment 3), middle aged plastids (segment 2), and mature plastids from the leaf tip (segment 1; Figure 3B). The Fs/Fm values were determined 6 h and 4 days after either a PSI-II or a PSII-I light shift (Figure 4A) in comparison to the respective controls. 6 h after a light shift only segment 1 revealed significant changes in Fs/Fm values. Segment 2 and 3 exhibited very weak Chl fluorescence without any recognizable differences between the growth light regimes. Four days after the light shift the Chl fluorescence in general was increased indicating the progress in the build-up of the photosynthetic machinery during the time range of observation. However, again segment 1 was the only segment displaying significant Fs/Fm fluorescence changes characteristically for a LTR.
FIGURE 4. Chl fluorescence and Chl a/b ratios in defined segments of maize seedlings acclimated in long-term to different artificial light qualities. Growth light conditions are given below the charts. Plants under PSI-light were shifted to PSII-light and vice versa and harvested 6 h of 4 days after the shift (indicated in the left margin) and compared to non-shifted controls. (A) Fs/Fm and (B) Chl a/b were determined from the same segments (indicated on top, compare Figure3). Data represent at least three independent biological replicates, standard deviation is given. A Student's t-test was performed to test significance of differences (*P < 0.05, **P < 0.01).
In the same experiments we analyzed the Chl a/b ratios of the respective samples (Figure 4B) which represents a second reporter for the LTR. After 6 h of acclimation it was found to be stable and comparable to the controls. After 4 days, however, the Chl a/b ratio of the PSI-II plants was significantly higher than that of PSI plants resembling that of PSII and WL plants. In contrast, PSII-I plants exhibited the opposite behavior with a lower Chl a/b ratio as in PSI plants. This pattern was found in all three segments analyzed indicating that even the younger leaf segments had begun to acclimate to the light condition although the Chl fluorescence measurements yet did not indicate this. A more detailed time course experiment indicated that the changes in Chl a/b ratios were largely finished after 2 days of acclimation (Figure A1 in Appendix) which is similar to the dynamics observed earlier in Arabidopsis (Wagner et al., 2008).
Redox-Controlled Molecular Processes in Photosynthetic Acclimation
Shifts between the PSI- and PSII-lights (and vice versa) induce reduction or oxidation of the plastoquinone pool (Wagner et al., 2008), respectively, which serves as a sensor for the balanced action of the two photosystems. In C3 plants, it controls the transcriptional regulation of the plastid gene psaA, targets the phosphorylation of the LHCII and the performance of state transitions. We tested all three processes in segment 1 of the leaf blades (Figure 5).
FIGURE 5. Redox-controlled processes in photosynthetic acclimation of maize seedlings. (A) Artificial growth lights were used as actinic light sources in state transition experiments to indicate the short-term redox effects on linear electron transport. Chl fluorescence given in arbitrary units was recorded using a PAM fluorometer. Measurements were done three times with leaves from three different biological samples. A typical trace is shown. On–off switches of the lights are given below the trace. Nomenclature of fluorescence parameters follows established conventions (see Materials and Methods). (B) Plastid protein extracts of maize seedlings grown under the different long-term light quality regimes as indicated in Section "Material and Methods" (indicated on bottom, compare Figures 3 and 4) were used to analyze the phosphorylation state of LHCII using a anti-phosphothreonine antibody. As loading control a corresponding experiment with the same membrane targeted to the small subunit of RubisCO (RbcS) is shown. (C) RNA isolates from the same samples isolated in parallel were used to determine the accumulation of psaA transcripts by northern analysis. As loading control a hybridization of the same membrane with a probe directed against the 18S rRNA is shown.
To demonstrate the effect of light sources on the redox state of the electron transport chain we used actinic lights in standard state transitions measurements (Haldrup et al., 2001). Chl fluorescence was detected using a pulse amplitude modulation (PAM) fluorometer (Figure 5A). The Chl fluorescence traces displayed changes characteristic for state transitions as observed earlier in C3 plants, i.e., a slow transient rise after a sudden drop when the far-red light was switched on and a more rapid transient decrease after a sharp rise when it was switched off again. These fluorescence transients are caused by the lateral migration of the LHCII between PSII and PSI and indicate that the time range necessary for state transitions in maize corresponded to that observed in Arabidopsis (Wagner et al., 2008).
Western analyses of thylakoid membrane phosphorylation state using an anti-phosphothreonine antibody (Figure 5B) indicated a strong phosphorylation of the LHCII proteins under PSII- and PSI-II light conditions while only weak phosphorylation signals could be detected in samples obtained from PSI- and PSII-I plants. This observation is consistent with the Chl fluorescence measurements and indicates the activity of a redox-sensitive kinase catalyzing state transitions via LHCII phosphorylation.
Northern analyses of total RNA preparations indicated a psaA transcript pool in PSI-light adapted plants which increased after a shift into PSII-light (Figure 5C). A similar transcript accumulation was observable in PSII-light grown plants and a down-regulation of psaA transcript amounts became apparent after a shift to PSI-light (Figure 5C).
In total all three parameters clearly demonstrated that redox-controlled acclimation responses occur in maize and that the seedlings perform a proper LTR at least in segment 1 under the artificial light quality regimes.
Recently, it could be demonstrated that the LTR includes targeted changes in the metabolic state of Arabidopsis and that this is reflected by a characteristic increase in the starch content of the leaves under PSII and PSI-II lights when compared to PSI and PSII-I conditions. The stn7 mutant was unable to perform this increase indicating that it is an redox-controlled process (Bräutigam et al., 2009). To test whether this regulation is also present in maize, seedlings were grown under the four light regimes and starch accumulation was determined in each segment (Figure 6). In contrast to Arabidopsis we could not observe any distinct pattern of starch accumulation in response to the light quality indicating that this regulation does not work in maize although the photosynthetic acclimation processes appeared to be fully functional. This result is consistent with the findings in plants collected from the field.
FIGURE 6. Starch accumulation in segments of maize seedlings grown in long-term under the different light quality regimes. Starch per fresh weight (FW) was determined from the three leaf segments (indicated on the right) as described before. Growth light regimes of the respective plant samples are indicated at the bottom of each panel. All determinations were done in triplicate, standard deviations are given.
Redox-Dependent Phosphorylation of LHCII in Maize Grown in the Field or Under Various Laboratory Conditions
In order to obtain a direct comparison of acclimation abilities of maize under field and laboratory conditions we determined the phosphorylation state of the LHCII using immunoblotting analyses with antibodies directed against phosphothreonine. We harvested leaf material in the field corresponding to the levels 1–3 (compare Figure 2) and isolated total leaf protein extracts. 20 μg protein were separated by SDS gel electrophoresis and subjected to immunoblot analysis (Figure 7). In parallel, we separated the same amounts of protein extracts isolated from 2 months old maize plants grown in growth chambers where they experienced mainly a light intensity gradient due to vertical growth without neighboring plants. In addition, we grew maize seedlings for 10 days directly under two different WL intensities (170 and 35 μE) as well as in our light quality regime. For the latter Arabidopsis plants grown in parallel were used as control. Redox-mediated LHCII phosphorylation induced by growth under PSII- and PSI-lights appeared to be comparable for Arabidopsis and maize seedlings (Figure 7, lanes 1, 2 and 12, 13) and resulted in a strong phosphorylation of LHCII under PSII-light and a weak phosphorylation under PSI-light, respectively. This result is in accordance with our earlier observations (Wagner et al., 2008, Figure 5). Field grown maize exhibited a phosphorylation state of LHCII at levels 2 and 3 (lanes 5, 6) being comparable to that of the PSI-light control (lane 13) while that of level 1 (lane 4) was clearly reduced. The latter observation corresponds to reports demonstrating that the LHCII kinase is inhibited at high light intensities by a thiol-dependent repression mechanism (Rintamaki et al., 2000). Interestingly, the phosphorylation signal occurring above the LHCII remained stable under all conditions. Because of size and regulation behavior of this band under lab conditions it is likely the D1 protein which requires experimental proof in the future. If this assumption is correct then the stable phosphorylation contrasts results from laboratory experiments with Arabidopsis which suggest that D1 phosphorylation is affected by light intensity (Tikkanen et al., 2008; compare also lanes 7–9) pointing to potential additional control mechanisms in field grown maize besides light-dependent redox control. Maize from growth chambers not subjected to dense planting displayed a regular increase in LHCII phosphorylation from the top level to the bottom (lanes 7–9) which can be easily explained by the light-intensity dependent repression of the kinases. The additional experiment with the 10-day-old maize seedlings grown under 170 and 35 μE WL (lanes 10 and 11) displaying high phosphorylation states under both conditions indicates that a significant high light repression of LHCII phosphorylation in maize likely starts beyond the 330 μE light intensity point (Lane 8). This is much higher as the corresponding point in Arabidopsis (about 150 μE, compare Bonardi et al., 2005), probably since maize is genotypically adapted to much higher light intensities for growth than Arabidopsis. In summary, we conclude that the principal LHCII phosphorylation events occurring in the field can be mimicked by our artificial light quality system. Phosphorylation of the putative D1 protein, however, appears to be potentially under a more complex control including additional influences besides the light environment and requires more complex physiological set-ups in order to study it.
FIGURE 7. Redox-dependent phosphorylation of LHCII in maize grown under field conditions or various laboratory conditions. Protein extracts of differentially grown plants were separated on a denaturing 10% SDS-PAGE, blotted to a PVDF membrane and subjected to an immunoblot analysis using an anti-phosphothreonine antibody (top panel). Signals were visualized by enhanced chemiluminescence. 20 μg total protein were loaded per lane. A parallel control gel was loaded with the same protein amounts from the same samples and subjected to Coomassie staining as loading control (bottom panel). Lanes 1 and 2: extracts of 10-day-old Arabidopsis seedlings grown under artificial PSII- and PSI-light serving as control. Lane 3: size marker. Lanes 4–6: Leaf protein extracts of 4 months old field grown maize (L1–L3 correspond to the respective height levels given in Figures 1 and 2). Lanes 7–9: leaf protein extracts of 2 months old maize plants grown in a growth chamber (GC) equipped with mercury vapor white light sources. Light intensity at the height of leaf harvest is given in μmol photons per m2 and s. Lanes 10 and 11: leaf protein extracts from 10-day-old maize plants grown in growth cabinets equipped with cool white light fluorescent stripe lamps (light intensity is given in μmol photons per m2 and s). Lanes 12 and 13: leaf protein extracts of 10-day-old maize plants grown under the same PSII- and PSI-light conditions as given for extracts from Arabidopsis seedlings presented in lanes 1 and 2. All plants used for experiments given in lanes 7–13 were grown in the same growth chamber being subjected to 60% relative humidity and 18–21°C.
Maize is one of the major crops in worldwide agriculture and, therefore, its ability to respond and acclimate to changing environmental conditions is of great interest, especially with regard to rising CO2 concentrations in the atmosphere (Prins et al., 2007). Recent analyses have demonstrated that the acclimation to CO2 is closely related to the water status and the developmental stage of the leaf investigated reflecting the multiple effects on photosynthesis (Prins et al., 2011). The efficiency of CO2 fixation, however, also largely depends on the energy provided by the light reaction which might create a limiting factor since it is well-known that maize fields possess a very high LAI. Our measurements of the light gradients within such a field confirm this and demonstrate that in a fully grown maize field approximately 70–80% of the biomass can perform photosynthesis only in a very sub-optimal way since the useful light wavelengths as driving force for the light reaction are very limited. This occurs even under optimal sun light conditions suggesting that there is a high potential for improvement of light usage efficiency in the same way as discussed for rice (Horton et al., 2001).
Recently, molecular studies investigated the molecular changes of the photosynthetic apparatus in maize in response to WL intensity gradients (Romanowska and Drozak, 2006; Romanowska et al., 2008). In the present study we focused on potential changes in photosynthetic functions in maize caused by long-term variations of incident light quality using artificial light sources exciting preferentially PSI or PSII. A similar approach has been performed earlier, however, with the main focus on the effectiveness of the phytochrome system (Eskins et al., 1986). A recent study explains photosynthetic acclimation to canopy density of tobacco and Arabidopsis as an interplay of photoreceptors and cytokinins, but the results also suggest a possible existence of alternative signaling pathways (Boonman et al., 2009). Our physiological set-up was originally established to analyze processes controlled by redox signals from photosynthetic electron transport in C3 plants. Our experiments presented here demonstrate that the system is able to induce a prominent short-term response also in the maize seedlings. Redox-controlled state transitions as measured by Chl fluorescence and corresponding changes in LHCII phosphorylation (Figure 5) were observable in a comparable way as reported earlier (Andrews et al., 1993). In addition, we could detect a pronounced LTR with corresponding changes in psaA transcript accumulation (Figure 5C) and characteristic changes in Fs/Fm and Chl a/b values (Figure 4). The LTR, however, was fully observable only in the top leaf segment of the maize seedlings indicating a developmental dependency of the LTR on the gradient of chloroplast biogenesis at least under the conditions of the present study (Figure 4). In contrast, in mature leaves of field grown plants the LTR as reported by respective changes of Fs/Fm and Chl a/b was readily detectable in all segments (Figure 2) supporting this conclusion. Our light measurements in the field confirm earlier results demonstrating that in dense plant population strong light quality gradients exist which are present even under sunny conditions (Figure 1). It can be concluded that fluctuations of illumination from outside are dampened in their absolute values by the top layers of the field and that within the field canopy the light environment is a rather stable low-intensity, far-red enriched light condition. Thus, leaves in the top level mainly require to perform high light acclimation using mechanisms such as non-photochemical quenching (NPQ). In contrast, for the largest portion of the biomass in the field a long-term acclimation to the non-saturating low light conditions appears to be more desirable. We could observe a distinct difference between the Fs/Fm and Chl a/b values for the top storey of the field when compared to middle or bottom storey indicating an adjustment of the photosynthetic apparatus to the respective illumination conditions (Figure 2). Leaf autonomous regulation of photosynthetic acclimation via redox signals from photosynthesis would provide an efficient and simple regulation principle allowing the individual plants to adapt to the strong gradient in light intensity on one hand and the parallel light quality gradient on the other. Thus, our physiological light system and the reporter for the LTR provide a useful test system not only for redox-controlled processes, but also for mimicking field conditions in dense canopies.
In Arabidopsis we observed a concomitant variation of the starch amount during the LTR which was reported to be a reporter for the metabolic changes occurring in the background of the light acclimation response (Bräutigam et al., 2009). Interestingly, this was not observable in maize, neither in the seedlings nor in the mature plants. In Arabidopsis, it was shown that starch synthesis is dependent on redox regulation of the entry enzyme ADP-glucose-pyrophosphorylase (AGPase) via the thioredoxin-like NtrC (Michalska et al., 2009). It is not clear yet whether the changes in starch accumulation during the LTR are also mediated by NtrC, however, it is a likely candidate. In maize obviously the regulatory link between LTR and starch accumulation has been uncoupled. A possible explanation would be the C4 syndrome of maize, since in C4 metabolism starch is mainly produced in the bundle sheath cells which do not contribute significantly to overall linear electron transport, thus lacking the basic requirement for photosynthetic redox signaling. However, it could be shown that even the agranal chloroplasts of the bundle sheath cells of maize contain a certain amount of PSII providing at least a limited capacity for redox signaling (Romanowska et al., 2008). BLAST searches done with the NtrC protein sequence of Arabidopsis identify an ortholog in maize with 92% positive amino acids strongly suggesting the existence of this key redox regulator in maize. Interestingly, starch accumulation in bottom leaves of the field grown plants is comparable or even slightly higher than in the top level leaves. Thus, starch accumulation appears to be uncoupled from the major site of primary photosynthesis implying a number of questions concerning the source–sink relationships in this plant which are an interesting topic for future work.
Regarding the high proportion of the field biomass performing a LTR the question arises for the beneficial effects of this acclimation response. In Arabidopsis, a clear negative impact on rosette growth and seed production could be demonstrated when the LTR is lacking (Wagner et al., 2008; Bräutigam et al., 2009). A recent study analyzed in detail the wavelengths dependency of photosynthetic quantum yield indicating significant positive effects for plants when performing a corresponding acclimation response (Hogewoning et al., 2012). Comparable experiments in maize require the isolation of corresponding LTR mutants which are currently under way. In addition, further field experiments are required to understand temporal and spatial occurrence of the LTR during the development of the plant stand. Young and small maize seedlings in the field perceive much more high light per total leaf area than the older plants which need to compete for light. Thus, the requirement for a LTR in the lower parts of the plants increases with the development of the plant stand. Understanding the molecular regulation of these processes will help to assess the potential of yield improvement in crop field via engineering of photosynthetic acclimation responses.
Materials and Methods
Field Grown Plant Material
For field analyses we characterized a plot in the middle of a fully grown maize field near Jena on the 9th of September 2009 at a time point when development of the corncobs has not yet started. The density of plants within the field was between 16 and 20 plants/m2. Light conditions in the maize field were measured at different indicated heights at noon under sunny and dry conditions and 22°C air temperature. Light intensity of PAR was determined with a Licor LI-250 (Heinz Walz GmbH, Effeltrich, Germany). Light spectra were determined with a spectroradiometer GER 1.500 (Geophysical and Environmental Research Corp., Millbrook, NY, USA). Reference data were obtained outside the field in a plant free area nearby which was also free of reflecting light from the fields around. Based on these measurements leaves were harvested at three different heights with characteristic light conditions. Only healthy, green leaves without any sign of beginning senescence or pathogen attack were chosen. Leaves were put in a light-proof box on ice and transported immediately to the lab. Chl fluorescence measurements were started within 30 min after harvest and after acclimation to room temperature. Material for Chl and starch determination was frozen in liquid nitrogen and kept at -80°C until further use.
Plant Material Grown in Climate Chambers
For analysis of young maize seedlings Z. mays (L.) var seeds were sown in pots on soil/vermiculite (4:1) and grown in growth chambers at 25°C, 80% humidity and a 16 h light/8 h dark cycle. Illumination was ~30 μmol photons/m2s provided by cool white fluorescence lamps. Seedlings were grown for 3–4 days until the first leaf had developed. Then pots were shifted for further 3 days to light cabinets equipped with light sources preferentially exciting PSI or PSII (PSI- or PSII-light, respectively). The spectral composition of the light sources have been described in detail earlier (Wagner et al., 2008). Both light sources (PSI- and PSII-light) had an comparable PAR of 20–30 μmol photons/m2s which was found to be sufficient to sustain photoautotrophic growth. After this pre-acclimation one half of the plants were shifted from PSII- to PSI-light (PSII-I plants) and from PSI- to PSII-light (PSI-II plants) while the other half remained in the light cabinets (PSI and PSII plants). Plants were then re-acclimated for further 4 days before they were harvested for analyses. In time course experiments samples were taken 6, 24, 48, and 96 h after the light shift. Maize plants grown under high WL conditions or for prolonged growth under WL were subjected to illumination from mercury vapor WL sources in the same growth chambers as the PSI- and PSII-light sources. Light intensities as indicated in the figures were obtained by adjusting the distance of the plants to the light sources placed on top of them. The respective PAR intensity was detected using the Licor LI-250 (see above).
Chlorophyll Fluorescence Measurements
In vivo Chl a fluorescence parameters were determined by video imaging under room temperature using a pulse amplitude-modulated FluorCam 700 MF device (Photon Systems Instruments, Brno, Czech Republic). Fs/Fm and Fv/Fm of plants acclimated to different light qualities were determined as described earlier (Wagner et al., 2008). We also compared basic photosynthetic parameters such as Fv/Fm detected by the Fluorcam with that of a PAM2000 (Heinz Walz GmbH, Effeltrich, Germany) and found the saturation flash of approximately 2000 μE sufficient to close all PSII centers for determination of Fm. State transitions were determined as described earlier (Lunde et al., 2000; Damkjaer et al., 2009) with intact single plant leaves using a PAM-101 fluorometer equipped with a PDA100 for digital data recording (WALZ, Effeltrich, Germany). PSII- and PSI-favoring light was provided by the growth light sources as described (Wagner et al., 2008). Saturation pulses were given for 800 ms by a Schott KL-1500 lamp at 5000 μmol photons/m2s. One representative of three replicates is shown.
Chlorophyll and Starch Determination
Plant material was ground in liquid nitrogen in a mortar. For determination of Chl concentration the pigments were extracted with 80% buffered acetone. Chl a and b concentrations were then spectroscopically determined according to Porra et al. (1989). Starch was determined using a standard colorimetric method (Magel, 1991) and the absorption coefficients described by Appenroth et al. (2010).
Northern Analysis
Preparation of total RNA from the top section of 10-day-old maize seedlings grown in the light quality system as described above was performed as established earlier (Fey et al., 2005b). 10 μg of total RNA were separated on a 1% denaturing agarose gel containing 7% formaldehyde, transferred to a nylon membrane and hybridized to a probe directed against a highly conserved sequence in the psaA gene (encoding the apoprotein of PSI) of Arabidopsis following established standard protocols (Sambrook et al., 1989).
Phosphorylation State of LHCII
Fresh leaf tissue of differentially grown plants was homogenized using a Waring® blender in ice cold homogenization buffer (HB; 50 mM HEPES-KOH, 0.33 M sorbitol, 10 mM KCl, 5 mM MgCl2, 10 mM NaF) at 4°C. Chloroplasts were recovered by centrifugation for 2 min at 2000 × g. Total chloroplast protein extracts containing 20 μg Chl were solubilized for 10 min in 40 μl buffer containing β-dodecyl-maltoside (BDM) at a final concentration of 1%. Proteins were denatured with 40 mM DTT, 2% SDS, 4% glycerol, and 150 mM Tris–HCL pH 6.8 for 10 min and subjected to SDS-PAGE using a 12% gel. The gel was either stained with colloidal Coomassie following manufacturer's recommendations (Roth, Karlsruhe, Germany) or used for western analysis. The stained gel was scanned with a LI-COR Odyssey infrared laser scanner at 700 nm to detect the dye fluorescence and protein loading was determined by quantification of fluorescence signals. Calculations of relative protein amounts were done with LI-COR Odyssey2.1 software (median background correction). For immunoblot analysis gels were electro-transferred to a polyvinylidene difluoride (PVDF) membrane (Roti-PVDF, Roth, Karlsruhe, Germany) and de-stained according to current protocols (Wittig et al., 2006). Phosphorylated proteins were detected with a polyclonal primary anti-phosphothreonine antibody (Cell Signaling Technology Inc., Danvers, MA, USA) and a fluorophore-coupled secondary antibody IRD800-anti-rabbit (Rockland Immunochemicals Inc., Gilbertsville, PA, USA). For testing phosphorylation state of field material the harvested leaves were frozen on dry ice directly at the site of harvest to avoid any change in phosphorylation during transportation. Total leaf protein extracts were isolated according to Pfalz et al. (2009) and used for immune-western analysis with anti-phosphothreonine antibodies as described above with the exception that the detection was performed with enhanced chemiluminescence (ECL). Samples collected from lab growth chamber-grown plants were treated in an identical manner to allow direct comparison.
Conflict of Interest Statement
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.
We thank N. L. Chrestensen Erfurter Samen- und Pflanzenzucht GmbH for providing seed material. This work was supported by the Deutsche Forschungsgemeinschaft.
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Pesaresi, P., Hertle, A., Pribil, M., Kleine, T., Wagner, R., Strissel, H., et al. (2009). Arabidopsis STN7 kinase provides a link between short- and long-term photosynthetic acclimation. Plant Cell 21, 2402–2423. doi: 10.1105/tpc.108.064964
Pfalz, J., Bayraktar, O. A., Prikryl, J., and Barkan, A. (2009). Site-specific binding of a PPR protein defines and stabilizes 5′ and 3′ mRNA termini in chloroplasts. EMBO J. 28, 2042–2052. doi: 10.1038/emboj.2009.121
Pfannschmidt, T., Nilsson, A., and Allen, J. F. (1999a). Photosynthetic control of chloroplast gene expression. Nature 397, 625–628. doi: 10.1038/17624
Pfannschmidt, T., Nilsson, A., Tullberg, A., Link, G., and Allen, J. F. (1999b). Direct transcriptional control of the chloroplast genes psbA and psaAB adjusts photosynthesis to light energy distribution in plants. IUBMB Life 48, 271–276.
Pfannschmidt, T., Schutze, K., Brost, M., and Oelmuller, R. (2001). A novel mechanism of nuclear photosynthesis gene regulation by redox signals from the chloroplast during photosystem stoichiometry adjustment. J. Biol. Chem. 276, 36125–36130. doi: 10.1074/jbc.M105701200
Pons, T., and de Jong-Van Berkel, Y. (2004). Species-specific variation in the importance of the spectral quality gradient in canopies as a signal for photosynthetic resource partitioning. Ann. Bot. 94, 725–732. doi: 10.1093/aob/mch197
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Walters, R. G., Rogers, J. J. M., Shephard, F., and Horton, P. (1999). Acclimation of Arabidopsis thaliana to the light environment: the role of photoreceptors. Planta 209, 517–527. doi: 10.1007/s004250050756
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Zhu, X.-G., Long, S. P., and Ort, D. R. (2010). Improving photosynthetic efficiency for greater yield. Annu. Rev. Plant Biol. 61, 235–261. doi: 10.1146/annurev-arplant-042809-112206
Kinetic changes of Chi a/b ratio in defined segments of maize seedlings acclimated to different light qualities. Growth light conditions are given in the right margins. Plants under PSI-light were shifted to PSII-light and vice versa and harvested at indicated time points after the shift and compared to non-shifted controls. Data represent at least three independent biological replicates, standard deviation is given.
Keywords: photosynthesis, redox regulation, light quality, light acclimation, maize fields
Citation: Hirth M, Dietzel L, Steiner S, Ludwig R, Weidenbach H, Pfalz J and Pfannschmidt T (2013) Photosynthetic acclimation responses of maize seedlings grown under artificial laboratory light gradients mimicking natural canopy conditions. Front. Plant Sci. 4:334. doi: 10.3389/fpls.2013.00334
Received: 27 June 2013; Paper pending published: 29 July 2013;
Accepted: 08 August 2013; Published online: 12 September 2013.
Edited by:Mohammad Mahdi Najafpour, Institute for Advanced Studies in Basic Sciences, Iran
Reviewed by:Suleyman I. Allakhverdiev, Russian Academy of Sciences, Russia
Helmut Kirchhoff, Washington State University, USA
Copyright © 2013 Hirth, Dietzel, Steiner, Ludwig, Weidenbach, Pfalz and Pfannschmidt. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.
*Correspondence: Thomas Pfannschmidt, Laboratoire de Physiologie Cellulaire & Végétale, Univ. Grenoble Alpes, 17 rue des Martyrs, F-38054 Grenoble, France e-mail: Thomas.Pfannschmidt@ujf-grenoble.
†Present address:Matthias Hirth, Institut für Allgemeine Botanik und Pflanzenphysiologie, Professur für Molekulare Botanik, Friedrich-Schiller-Universität Jena, Dornburger Straße 159, Jena 07743, Germany;
Lars Dietzel, Institut für Molekulare Biowissenschaften, Pflanzliche Zellphysiologie, Biozentrum Goethe-Universität Frankfurt, Max-von-Laue-Straße 9, Frankfurt am Main 60438, Germany;
Sebastian Steiner, Klein Wanzlebener Saatzucht Saat AG, Grimsehlstraße 31, Einbeck 37574, Germany;
Robert Ludwig, Institut für Diagnostische und Interventionelle Radiologie I — AG Experimentelle Radiologie, Universitätsklinikum Jena — Friedrich-Schiller Universität Jena, Erlanger Allee 101, Jena 07747, Germany;
Thomas Pfannschmidt, Laboratoire de Physiologie Cellulaire & Végétale, Univ. Grenoble Alpes, 17 rue des Martyrs, Grenoble F-38054, France


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Key Investment Considerations
Reason #1:
Big Yield (9%)
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Cheadley Wheaps
The Manual Masters talk Octopath Traveler some more, argue about Naruto and come up with ways to make NPCs "stick" in this Gruffin Tumble adventure.
Sponsored by Mr. Technimatter's Brain Splatter
Questions Answered:
Do you have any suggestions for how to build a character that really makes a lasting impression on the players?
I was just wondering if you had any tips or tricks for a Naruto based campaign or adapting a campaign to a different universe.
Loot Table:
The Never Known
To-Do Wrist
Monster: Gruffin
Relevant Links:
Mutants and Masterminds


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Power BI Development & Consulting
Combining the 4 phases of data: Reporting, Analysing, Monitoring & Predicting to enable better decision making across all all layers of an organisation
Combining the 4 phases of data: Reporting, Analysing, Monitoring & Predicting to enable better decision making across all all layers of an organisation
Icreon works with analytics and data leaders, typically on the Marketing and Operations side, to help get the most out of their information. We do this by being a Microsoft Gold Partner and aligning ourselves on one of the strongest Business Intelligence and data analytics platforms in the marketplace, PowerBI.
In addition to work on data and analytics strategies with our clients, we work directly with operational stakeholders, such as data scientists, machine learning researchers, and analysts to craft out the best ways to implement a PowerBI implementation from start to finish. This includes evaluating how to aggregate data, what the most pertinent statistical models are, and the best ways to visualise them for varying stakeholders.
Talk to a PowerBI specialist.
Creating clear problem statements for the business that we can use as a north star to define our implementation strategy
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Interpreting the results of data collection by using advanced models to find trends within your business
Putting new business practices into place to take advantage of the prior data analysis and repeating the loop


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At Discover, be part of a culture where diversity, teamwork and collaboration reign. Join a company that is just as employee-focused as it is on its customers and is consistently awarded for both. We're all about people, and our employees are why Discover is a great place to work. Be the reason we help millions of consumers build a brighter financial future and achieve yours along the way with a rewarding career.
Accountable for proactively and holistically leading activities, consultation, and documentation that guide the application development community through a portfolio of technology components and varying deployment patterns. Works collaboratively to design, document, and request delivery of multiple concurrent, highly technical, and financially visible complex projects. These projects will leverage multi-platform and multi-site/zone infrastructure environments.
The Cloud Solution Architect is an exciting role that has one foot planted in cloud security and the other in DevOps / Automation. This is a fast-paced role, well suited for someone who enjoys keeping their existing technical skills honed and developing new ones; so you can make strong contributions to deep architecture discussions. As a member of the Solutions Architecture team, this individual will be responsible for requirements gathering, design, and high-level socialization of complex infrastructure solutions needed to solve an IT/business problem. The ideal candidate must be self-motivated with a proven track record in software/technology consulting. The ability to connect technology with measurable business value is critical to a solutions architect. You should also have a demonstrated ability to think strategically about business, products, and technical challenges.
Your broad responsibilities will include: owning the technical engagement and ultimate success around specific implementation projects; defining implementation architectures, responding to questions, developing a deep expertise in cloud technologies as well as broad know-how around how applications and services are constructed using the cloud (AWS platform). The scope of this role includes heavy participation in technical and non-technical project meetings, development of presentation quality technical documentation, creative conflict and issue resolution, and communication of design criteria and direction across all levels of IT management. A candidate must have high-level communication skills and be comfortable working with internal customers, management, technical staff, and external vendors to assess needs, requirements, and design standard/custom infrastructure solutions as necessary.
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General Malaise Monday
I've been listening to Mature Themes by Ariel Pink's Haunted Grafitti a lot lately. Makes for good background noise when I'm doing things that require serious concentration, like googling the "cutest picture on the internet" and getting results like this:
Have a good week y'all!
Follow me over here, ya dig?


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Unplanned Adventures: The First Couple Days
As we all know, plans are best done in our heads and left out of the hands of reality, or as the saying goes, the quickest way to make God laugh is to tell Him our plans. Everything was going smoothly the first day (our travel day), until we got to Badlands National Park. The plan was to eat at a restaurant located there and drive through the badlands, taking in its natural allure until we reached our campsite. Unfortunately, we realized too late that we needed gas in the cars and the only gasoline station near us was back out the same way we drove in. This would take some extra time and also meant that night would quickly be approaching. There was simply not enough time to take the 40+ minutes drive through the Badlands unless we desired to set up camp in the dark. It was educators Stephen and Angelyn who stepped up and volunteered to drive the quicker way, taking the interstate around most of the park. While they were doing that, everyone else took the longer, but maybe more harrowing drive across these untouched lands in the twilight hours. In the end everyone made it safely to the campsite and we still ended up setting up a couple things in the dark. Even our more diligent plans seemed to fail…and of course we forgot a rain tarp for one of the tents!
This is not to say that any of us were discouraged by these unexpected course corrections. In fact, they added a bit of thrill to the experience. We woke up in the morning to a stunning sky and roaming buffalo next to the campsite. Educator John Eilen even took some time to commune with the local prairie dog community (he fit right in, as the prairie dog is his spirit animal). We took in the sights, which made for a great early morning, and set off for Mass in Rapid City at the Cathedral of Our Lady of Perpetual Help. Mass was refreshing, as it always seems to be, and we had some extra time to view the church and pray both before and after. The next stop was Mount Rushmore, so with quick jaunt from Rapid City, we viewed the impressive mountain sculptures, took in the history, and had a nice little hike. At this point, our second change of plans became inevitable. We intended to hike around Sylvan Lake that afternoon but the ever present enemy, fatigue, had set in on the party; so, it was decided we would take some time at the gift shop and then head over to the campsite early to start cooking supper.
The final installment coming soon!


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Ljubljana | The Ultimate City Guide/
In the past few years Slovenia has started to slip into travel bucket lists around the world. It's capital city, Ljubljana (Lub-lee-ann-uh) is a perfect European city break with fusions of Central European and Mediterranean influences, chilled vibes, beautiful wine and beer, the cutest Cafes and delicious food.
In this guide, you will learn how to get around, places to stay, top things to do, best restaurants as well as shopping recommendations.
* * Disclaimer: I have added affiliate links to this post, which means if you click and buy I will earn commission. It helps me keep Earth Lists going, so thanks in advance.
Currency: Euro
Ljubljana Airport:
It's 26km away from Ljubljana and roughly takes 25 minutes. Taxi's cost around €35 one way. Bus 28 departs every hour from the airport to the city centre, and costs €4.10. If you are hiring a car, walk out of the main terminal and directly across to the building in front of you.
Getting Around:
Ljubljana is a small city and flat. Walking is the best way to see the city, so make sure you pack a comfy pair of trainers. However, like Copenhagen, the city is bike friendly with designated lanes.
Use Bicike(LJ), these 38 bicycle stations are dotted around the city. You have to pay a registration fee of €1. Six hours will cost you about €15, however, the first hour is free, so if you are savvy you can ride to you location dock the bike and pay no more than €1. Bargain.
If you are planning on being in Ljubljana for more than 2 days I would strongly suggest you hire a car and visit Lake Bled, Piran, Trieste, Triglav National Park or Vintgar Gorge.
Slovenia is quite cheap in comparison to its neighbouring countries. On average I paid €3 - €4 for a glass of Wine or beer. A meal for four with starters, mains and drinks was no more than €75.
Where to Stay in Ljubljana
Ljubljana is super small compared to other cities so accommodation options are limited. I stayed in an Airbnb, a 10 minute walk out of the city because the hotels were full the week we decided to go. However, there are a few boutique hotels doted around and I took note of a few that I liked the look of.
Firstly, there are two on Gornji trg, *Allegro and *Lesar, both are boutique hotels and are situated on the cutest street in the old town. One of these would be my first option. If not, check out *Hotel Cubo which is on the other side of the river but has outstanding reviews. Prices vary but you will be paying around €120 per room per night.
Take a look at Booking.com deal finder to get your search underway!
Things to do in Ljubljana
As Ljubljana is so small, I wouldn't say there is tons of sightseeing to do, which I love as sometimes we get so preoccupied with list ticking we don't get to explore. There are lots of pretty squares, streets, architecture and museums to check out though. Here are some of the top things you can do.
Prešeren Square
This is the main square in Ljubljana and what a beauty she is. Here you will find the triple bridge and the gateway between the Old Town and Shopping area.
Galerija Emporium
High End Luxury Department Store with one hell of an impressive Art Nouveau staircase. Even if you don't fancy splashing out on a fancy handbag or manbag, it's worth a look for the staircase alone.
Ljubljana Castle
The Castle is literally on the hill (cue Ed Sheeran song) overlooking the city. Take the Glass Funicular Up, a single ticket is €2.20. Castle entry with Funicular is €10. Now I was a little underwhelmed when I got to the top, so for me it's not high on my priority list, however, if you like Funiculars and exploring castles, eat your heart out.
Dragon Bridge
Ljubljana's symbol is a dragon, they are everywhere. Plecnik designed the bridge and quite a lot of Ljubljana. It's a lovely stroll down to and is on the more quieter side of the city.
Stari Trg and Mestni Trg
Two of my favourite cobbled streets. Full of restaurants, Boutique shops, Bars and stunning architecture.
Odprta Kuhna ( Food Market) – Every Friday Spring and Summer.
Luckily, I was there to enjoy their 100th Celebratory party. It's a must to eat here if you are visiting on a Friday. Local Restaurants have pop up stalls and there are many international cuisines. Dishes vary from €4-6 Euros, wine is €2.50 and beer €3.50. Grab a few different plates and share. You can sit by the River and soak up the chilled vibes.
Walk along the River Ljubljanica
Strolling alongside the River is a must day and night.The vibes completely change at night, when the city becomes more, well, lively. The sound of music and chatter is the air, soak it all up with a glass of Movia Merlot in one of the many riverside bars. Cafe Romeo and Tozd were particularly ambiant at dusk.
Tivoli Park
Unfortunately, I didn't get the chance to visit Tivoli Park as I had a lot to cram in. However, if the weather is nice and you have a spare couple of hours, do have a wander around these impressive gardens.
Visit Lake Bled and Triglav National Park from Ljubljana
A day trip to one of the most beautiful lakes and National Parks in the World should not be missed. Read how to get to Lake Bled from Ljubljana
If you're looking for a guided tour, check out Get Your Guide below for the best selling options.
Where to eat in Ljubljana
Quite the culinary city, Ljubljana is a delight for serving up mouth-watering cuisine. From fresh pasta to Goulash to stacked burgers, there is an array of international influences to choose from. Some of my personal favourites:
Julija | Stari trg 9
One of my favourite meals in any country, no wonder it was packed out most nights. Make sure you reserve a table here. Surprisingly, inexpensive considering just how delicious the food was - we paid €72 for 4 people (mains, wine and beer).
Verace | Streliška 22
The Best Pizza in town. Recommended by our host, I am so glad we decided to stop here for lunch. Affordable and tasty with a huge selection of pizza's to choose from. Pizza's range from €5 - €9.50. A little walk out of the centre but is worth it. Be warned, it is closed on a Sunday and Monday.
AS Aperitivo | Čopova ulica 5a
I stumbled upon AS Aperitivo and I am so glad, this dinner was incredible. I can only describe this restaurant as Italian LUXE, the interior is stunning. The service was second to none and the food was delicious. I had Tortellini and Parma Ham and washed it down with a local Pinot Grigio...oh take me back. Prices weren't bad at all, averaging €15 for a main and drink.
Cafés in Ljubljana
Lolita | Cankarjevo nabrežje 1
Hands down the prettiest café I've been to. With a smorgasbord of tasty cakes and chocolates to sample, you will be spoilt for choice. We couldn't help ourselves and had four cakes as well as a bag of chocolates, which came to €25. The waiter knows his stuff so ask him to point you to your perfect treat. Your Welcome!
Le Petit Café | Trg francoske revolucije 4
Simply, the best breakfast I had the whole trip. This French - styled cafe has a wide range of breakfast items at affordable prices, we paid €34 for four people including two rounds of hot drinks...bargain.
Pritličje | Mestni trg 2
It's an Interior Designers' dream here. Full of greens, hanging plants and ultra modern, with a small comic book stall thrown in for good measure. Free Wifi and the coffee is Damn Fine. It's located on the main street in Old Town so make sure you stop here.
Shopping in Ljubljana
Ljubljana is full of the quaintest and cutest boutique shops as well as main stream stores. As mentioned previously, if you want High End luxury make sure you visit Galerija Emporium or head to BTC and City Park Shopping Centres.
GUD | Stari trg 4
There was so many amazing things I wanted to buy. GUD reminds me of the boutique stores in Copenhagen, filled with Scandi Goodness. I especially love the Ljubljana prints, check out the one I brought below, accurate, cute and now on my wall.
Smile Concept Store | Mestni trg 6
Don't forget to Smile, is the tag line for this store and when you step through the door you are met with CROSS THIS LINE WITH A SMILE graffiti on the floor. Full of cool and quirky gifts and even a selfie wall, make sure you stop off and be infected by the Smile Concept :)
Babushka | Stari trg 18
Not only does it have the coolest name (we love you Kate Bush), it has a wide range of the cutest home ware products and gifts. From Jewellery, to prints, to collapsible shelves and Air Plants, there is something for everyone.
Slovenian Wine Tasting in Ljubljana
Movia | Mestni trg 2
Without a doubt, go to Movia and get tasting Slovenian Wine, in my opinion, some of the best wine I've tasted. The Wine connoisseur here knew everything about each wine and took us on an informative and delicious journey through the regions of Whites and Reds. We also ordered a glass of Orange Wine (never knew Orange wine existed!) it's Red grapes treated like white. Prices were great too, as you can see below €10 - €16 for four glasses. We paid €18 per person for a sampler of White, Red, glass of orange and nibbles...I love Slovenia.


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Jaime D w/ Kim Leduc defeated Anna Minoushka
Deanna Conda defeated Jaime D w/ Kim Leduc to win the 2008 W.I.L.D. Tournament and the W.I.L.D Championship Title
Non-tournament Matches:
LuFisto defeated Phil Latio in an Intergender Hardcore Match to retain the ALF Championship
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