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Paleolithic diets include a moderate amount of protein, and have gained a lot of attention recently. The theory behind this dietary pattern is that our genetic background has not evolved to meet our modern lifestyle of calorically dense convenience foods and limited activity, and that returning to a hunter-gatherer way of eating will work better with human physiology. This has been studied in a few small trials, and it does seem beneficial for people with type 2 diabetes.
As a renal dietitian my focus can't be solely on diabetes. Although a very large percentage of our patients with chronic kidney disease are here due to unmanaged blood sugar control, that is just one of our problems. I have to prioritize my counseling in other ways – the most important being control of potassium, then sodium (fluid), protein and phosphorus.
A terrific rule to follow is to use a luncheon size plate rather than a dinner plate. This is an easy way to guide your portion sizes without having to think so much about it. Enjoy a glass of red wine or similar drinks only occasionally since your body treats alcohol more like a fat than a carb when it comes to calories. Eat a sound breakfast and try to eat a bigger lunch so the bulk of your calories are consumed by the midafternoon as a way to keep your blood glucose level in a healthy range, and lessen the chance of undesirable weight gain.1
If you like, I can help show you Medicare coverage options that may fit your health goals. To learn more, click the links below to set up a phone appointment or have me send you some personalized plan options by email. If you'd prefer to browse on your own, you can do that too: just click the Compare Plans button on this page to view coverage options in your zip code.
Diabetic's dietary practices are mainly influenced by cultural backgrounds. Concerning each of the dimensions of dietary practices, there were significant positive relationships between knowledge regarding diabetic diet and dietary practices. Knowledge was a salient factor related to dietary behaviors control.46 Moreover, patients' knowledge on a recommended diet indicates their understanding of dietary guidelines which influenced their food selection and eating patterns.47 The association between dietary knowledge and dietary practices among T2DM patients in the previous studies were inconsistent. Another study revealed that there was no relationship between dietary knowledge and compliance of dietary practices.58 On the other hand, the same study found that a high dietary knowledge score was associated with following dietary recommendations and knowledgeable patients performed self-management activities in a better way. Dietary knowledge significantly influences dietary practices. In Indonesia, a study was conducted to measure dietary practices among diabetic patients, which elaborated that the Indonesian people, preferred to consume high-fat foods which lead to an increased risk of CVD.59 The trend of skipping breakfast has dramatically increased over the past 10 years in children, adolescents, and adults.60,61 There is increasing evidence that skipping breakfast is related with overweight and other health issues.62 In addition, frequent eating or snacking may also increase the body weight and risk of metabolic diseases.63,64 Rimm65 demarcated western and prudent dietary patterns. The prudent dietary pattern was characterized by increased consumption of fish, poultry, various vegetables and fruits whereas; the western dietary pattern was characterized by an increased consumption of processed and red meat, chips, dairy products, refined grains, and sweets and desserts. These patterns were previously associated with T2DM risk. The glycemic index is an indicator of the postprandial blood glucose response to food per gram of carbohydrate compared with a reference food such as white bread or glucose. Hence, the glycemic load represents both the quality and quantity of the carbohydrates consumed.66-69 Another study conducted in Lebanon demonstrated direct correlation of the refined grains and desserts and fast food patterns with T2DM, however, in the same study an inverse correlation was observed between the traditional food pattern and T2DM among Lebanese adults.70
Foods might sometimes appear to be packaged into individual serving sizes even though they contain two or more servings per package. To determine that, look at "serving size" and "servings per container" at the top of any food label. For example, if a serving size is 1 and there are 2 servings per container, you will need to double all of the nutrient values on the label in order to get a clear picture of the value of the entire container.
One of the biggest tips I discuss with my clients who are trying to manage their Diabetes is to focus on adding more fiber to their diet. We first start by adding fruits and veggies to the meals that they are already consuming. I find adding to the diet is usually an easier approach for most. Another tip is to focus on adding more whole foods, like apples with the skin versus juices or baked sweet potatoes versus the instant varieties. Along with more fruits and veggies, clients can also add fiber in the forms of nuts and seeds to meals or snacks to help with satiety and blood sugar control. One of the key points I stress to any client is to focus the balance and take it one step at a time. I don't expect anyone to do a complete 360 on their food intake since it takes time to build new (and healthy) habits that will last long-term.
The types of fats in your diet can also affect the development of diabetes. Good fats, such as the polyunsaturated fats found in liquid vegetable oils, nuts, and seeds can help ward off type 2 diabetes. (39) Trans fats do just the opposite. (8, 40) These bad fats are found in many margarines, packaged baked goods, fried foods in most fast-food restaurants, and any product that lists "partially hydrogenated vegetable oil" on the label. Eating polyunsaturated fats from fish—also known as "long chain omega 3" or "marine omega 3" fats—does not protect against diabetes, even though there is much evidence that these marine omega 3 fats help prevent heart disease. (41) If you already have diabetes, eating fish can help protect you against a heart attack or dying from heart disease. (42)
Like refined grains, sugary beverages have a high glycemic load, and drinking more of this sugary stuff is associated with increased risk of diabetes. In the Nurses' Health Study II, women who drank one or more sugar-sweetened beverages per day had an 83 percent higher risk of type 2 diabetes, compared to women who drank less than one sugar-sweetened beverage per month. (26)
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Eventually, even though the pancreas is working at its best to produce more and more insulin, the body tissues (for example, muscle and fat cells) do not respond and become insensitive to the insulin. At this point, overt diabetes occurs as the body is no longer able to effectively use its insulin to maintain normal blood sugar levels. Over time, these high levels of sugar result in the complications we see all too often in patients with diabetes.
Whole grains don't contain a magical nutrient that fights diabetes and improves health. It's the entire package—elements intact and working together—that's important. The bran and fiber in whole grains make it more difficult for digestive enzymes to break down the starches into glucose. This leads to lower, slower increases in blood sugar and insulin, and a lower glycemic index. As a result, they stress the body's insulin-making machinery less, and so may help prevent type 2 diabetes. (22) Whole grains are also rich in essential vitamins, minerals, and phytochemicals that may help reduce the risk of diabetes.
How do sugary drinks lead to this increased risk? Weight gain may explain the link: In both the Nurses' Health Study II and the Black Women's Health Study, women who increased their consumption of sugary drinks gained more weight than women who cut back on sugary drinks. (26, 28) Several studies show that children and adults who drink soda or other sugar-sweetened beverages are more likely to gain weight than those who don't, (28, 30) and that switching from these to water or unsweetened beverages can reduce weight. (31) Even so, however, weight gain caused by sugary drinks may not completely explain the increased diabetes risk. There is mounting evidence that sugary drinks contribute to chronic inflammation, high triglycerides, decreased "good" (HDL) cholesterol, and increased insulin resistance, all of which are risk factors for diabetes. (32)

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Jun 13, 2019
We discuss the ongoing pace of play and length of game problems for MLB, automation of calling balls and strikes, Rougned Odor, Dallas Keuchel, Craig Kimbrel, and player pay
May 8, 2019
We consider the NHL's strengths and weaknesses, the revival of scoring in the NBA over the past 20 years, and the cities on MLB's short list for possible expansion sites.
Apr 10, 2019
We examine how the dominance of some superstars like Tiger Woods and some teams boost the popularity of a sport rather than diminishing it. We at the best of 7 playoff format for the NBA, and possible fixes for the high rate of strikeouts in MLB. We finish with a little bit on Bruce Pearl.
Mar 6, 2019
We address whether the new contracts for Manny Machado and Bryce Harper make sense, the positives and negatives of NCAA men's basketball, and differences in "fan experiences" particularly for teams we have followed such as the Mets or Kentucky basketball.
Feb 8, 2019
We consider the decline in the 2019 Super Bowl viewership in context with the uptick in viewers for the NFL regular season and playoffs. We examine pace of play proposals for MLB along with how Atlanta has become a soccer center in the U.S.


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This content is strictly the opinion of Dr. Josh Axe and is for informational and educational purposes only. It is not intended to provide medical advice or to take the place of medical advice or treatment from a personal physician. All readers/viewers of this content are advised to consult their doctors or qualified health professionals regarding specific health questions. Neither Dr. Axe nor the publisher of this content takes responsibility for possible health consequences of any person or persons reading or following the information in this educational content. All viewers of this content, especially those taking prescription or over-the-counter medications, should consult their physicians before beginning any nutrition, supplement or lifestyle program.
"The struggle is a normal, necessary part of the process. Changing your food is hard. Changing your habits is even harder. Changing your relationship with food is the hardest part of all. The process requires struggle—it's how you know you're growing—but don't make it harder than it has to be! There is no such thing as the 'perfect Whole30,' so if your beef isn't grass-fed or your travel meal doesn't look exactly like our meal template, don't sweat it. Your only job is to stick to the Whole30 rules for 30 days, and some days, you'll have to let good enough be good enough. When you do struggle, remember why you took on the program in the first place, and don't be overwhelmed by the big picture—just focus on the next day, or the next meal. Don't be afraid to ask for help when you need it, and high-five yourself for the victories you're achieving every day you're on the program, no matter how small. Even tiny progress is progress."
Smoothies are generally discouraged on Whole30. However, if you must have one, choose a recipe with no added sweeteners and low-sugar fruits. This smoothie is more scoopable with a spoon, so you can slow down and savor it (important on Whole30!). The only sweet flavor comes from fresh raspberries — plus, this packs your bowl with fats from avocado and coconut milk.
This spring, Dr Cordain did an interview answering ten questions about the basics of The Paleo Diet®. To start your New Years out right, we wanted to share his answers with you. We hope you enjoy! - The Paleo Diet Team 1. The Paleo diet can be traced to a 1975 book by Walter Voegtlin, but, correct me if I'm wrong, you are responsible for bringing this diet to popularity in your 2002 book "The Paleo Diet." Can you me about your...
If you were to eat an unlimited amount of red meat (which the paleo diet technically allows), you may see your heart health suffer. While experts applaud the omission of packaged and processed foods like cake, cookies, chips, and candy — which are well known to be bad for your ticker — they're not crazy about the fact that paleo doesn't allow you to eat whole grains, legumes, and most dairy. Whole grains in particular have been linked with better cholesterol levels, as well as a reduced risk of stroke, obesity, and type 2 diabetes. (13) These are all comorbidities of heart disease. (14)
The Paleo diet can be traced back to gastroenterologist Walter Voegtlin, who brought up the idea of eating like our ancestors in 1975 in his book "The Stone Age Diet." A decade later, researchers Melvin Konner and Stanley Boyd Eaton published a paper on the paleo diet in the New England Journal of Medicine, which is considered one of the foundations of the Paleo diet as we know it today.
Update: So I just finished my first whole30 round and I have to say that I'm super excited by the results. When I hopped on the scale today, I noticed that I had lost 26.6 pounds. I then measured myself and noted that I lost a total of 31.5 inches (-5" from chest, -2" from right arm, -3.5" from left arm, -8" from waist, -5" from hips, -4" from right thigh, and -4" from left thigh). I am really excited by my results!
Another 2014 study in the European Journal of Clinical Nutrition compared the effects of the Paleo plan to those of a standard low-fat diet on 70 obese, postmenopausal women. After six months, the Paleo group lost 14 pounds on average, while the other group lost nearly 6 pounds. After a year, the Paleo group had lost 19 pounds on average, and the low-fat dieters had dropped 10 pounds. A year later, both groups had regained some weight: The Paleo group was still down 10 pounds, while the low-fat group had dropped an average of more than 6 pounds.
Some specific foods that fall under this rule include: pancakes, waffles, bread, tortillas, biscuits, muffins, cupcakes, cookies, brownies, pizza crust, alternative flour pastas, cereal, or ice cream. No commercially-prepared chips (potato, tortilla, plantain, etc.) or French fries either. However, this list is not limited strictly to these items—there may be other foods that you find are not psychologically healthy for your Whole30. Use your best judgment with those foods that aren't on this list, but that you suspect are not helping you change your habits or break those cravings. Our mantra: When in doubt, leave it out. It's only 30 days.
An Interview with Ward Nicholson now has three parts on the web. Good overview of man's diet over the past 65 million years. Long but highly recommended reading. First published in Chet Day's "Health & Beyond" newsletter. Now part of a very comprehensive Beyond Vegetarianism site. Every argument that your vegetarian friends use to avoid meat for health reasons is debunked here.
Healthy, delicious, and simple, the Paleo Diet is the diet you were designed to eat. If you want to lose weight—up to seventy-five pounds in six months—or if you want to attain optimal health, The Paleo Diet will change your life now. Dr. Loren Cordain, the world's leading expert on Paleolithic nutrition, demonstrates how by eating all the lean meats and fish, fresh fruits, and nonstarchy vegetables you want, you can lose weight and prevent and treat heart disease, cancer, osteoporosis, metabolic syndrome, and many other illnesses. Incorporating all the latest breakthroughs in Paleo nutrition research, this new edition of the bestselling The Paleo Diet includes six weeks of meal plans to get you started on the Paleo path to weight loss, weight control, increased energy, and lifelong health.
At CFIH, we believe that results are rewarded to those who work hard for them. We provide the tools and together we will work to accomplish your goals. We believe that it's not just our commitment to fitness and clean eating that helps pave the way to success, but also our community which holds all of the key pieces together. Fall in love with our culture and join a community that is filled with individuals who live lives of integrity, who exude a confident but humble demeanor, and who are committed to creating positive habits to last a lifetime! Join us and become a happier, healthier, and more fulfilled YOU!
Thank you CFA coaches for always scaling the WOD for my medical needs, but still making the WOD fun and challenging. I may not be able to lift heavy or ever be a part of the CF games, but I do feel like a part of CFA family and with that I could never go back to a gym. It's been a privilege meeting and working out with great people. I feel honored to be selected CFA Athlete of the Month. CROSSFIT PROUD!!!
TBK Fitness Program by Tamir Katz shows how to achieve fitness through a healthy, natural hunter-gatherer diet along with a comprehensive exercise program with over 60 different bodyweight exercises of varying difficulty targeting all of the muscles in the body. Also included is a detailed discussion of nutrition and the diseases of civilization based on scientific research, information on stress management and preventive medicine, recommendations on vitamin and supplement use, tips on how to make your fitness program succeed where others have failed, tips on food shopping and preparation, sample meals, and more. The Amazon reviews average to 4+ stars.
The Open, introduced in 2011 and so-called because participation is open to anyone,[23] is held over five weeks in February–March; a new workout is released on each Thursday night (Pacific Time) and competitors complete the workout and submit their scores online by Monday evening, with either a video or validation by a CrossFit affiliate. Since 2013, Open workout announcements have been broadcast live, and featured two or more past CrossFit Games athletes competing head-to-head immediately following the workout description. Beginning with the 2019 Games, the top athlete from each country and the top 20 overall Open finishers qualify directly to the Games. The Open is also used for seeding purposes at the Games even if an athlete qualified through the sanctioned events; if an athlete qualifies through a sanctioned event but does not do the Open, they will be seeded at the bottom.
A metaresearch group at Tilburg University in the Netherlands investigates scientists' research methods and operations, scrutinizing questionable practices such as selective reporting of statistical tests and data massaging. The group instead advocates for practices such as preregistering studies and making experimental data immediately available for open external review and verification. Its members are optimistic that "the perverse incentives of careerist academia, to hoard data and sacrifice rigor for headline-generating findings, will ultimately be fixed."


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Awesome experience with Charlie's Rooter Service! Our outside area drains were overflowing when it rained and about to flood toward the house. I called Charlie's and they had Luis come over within an hour. Luis was super friendly and ready to help when he arrived. He figured out the problem and snaked the street drain and pulled out a huge clump of roots that were clogging it up! But there was still some backup flooding on the side that he was able to isolate to get rid of the mud and debris that were clogging that up. Before leaving, Luis even gave suggestions on how to flush it out periodically to keep things flowing smoothly. Luis is the best and I would definitely use him for future plumbing problems, highly recommended!
Not all plumbing companies can guarantee up-front pricing, superior workmanship and parts, licensed and insured plumbing contractors, and courteous support from uniformed professionals. Whether you are in need of drain cleaning or would like to schedule a plumbing inspection for your home, Mr. Rooter's plumbing services can handle it all! Our skilled plumbers are knowledgeable and equipped with the latest tools needed to achieve optimal results. Give us a call at (855) 982-2028, or request a job estimate online.
Howard County Plumbing Service is a family owned and operated plumbing service company headquartered in Eldersburg, Maryland. We are a full service plumbing company with 25 years of experience and provide 24 hour Emergency Service Monday through Friday after 5pm. We limit our emergency service availability as we believe in the importance of quality time spent with our families. Our team is dedicated to providing our customers with prompt, courteous and professional service. Our goal is to build ongoing relationships with our customers by providing top notch service with the utmost respect for our customers and their homes/businesses. We aren't satisfied until our customers are satisfied.
Alexa is an experienced lifestyle and news writer, currently working with Reader's Digest, Shape Magazine, and various other publications. She loves writing about her travels, health, wellness, home decor, food and drink, fashion, beauty, and scientific news. Follow her traveling adventures on Instagram: @living_by_lex, send her a message: [email protected], and check out her website: livingbylex.com
A rooter service is a type of plumbing repair – it is any method of drain cleaning or pipe repair. Typically, it is when a plumbing service company uses a range of different tools to perform water pipe repair. It involves running water through your drains, using tools that they snake through your drains to unclog them and sometimes taking parts of the water pipe system apart so that they can take out some of the clogs manually. The original culprit of clogged drains was tree roots – hence the name ROOTer service, but now it is used to address all kinds of stubborn sewer line and drainpipe clogs that affect the function of your plumbing.
Mr. Rooter® Plumbing is the go-to plumber for homeowners across the country. Why? Our plumbing contractors provide courteous, friendly, affordable, and effective residential plumbing services. We take our mission to exceed our customers' expectations seriously! From on-time arrivals to licensed and certified plumbing experts, we make sure homeowners are getting the very best plumbing services. You can have total peace of mind and confidence when you call on us! We stand behind our work and go the extra mile to get the job done quickly so you can get back to enjoying your home.
Wall thickness does not affect pipe or tubing size.[20] 1/2" L copper has the same outer diameter as 1/2" K or M copper. The same applies to pipe schedules. As a result, a slight increase in pressure losses is realized due to a decrease in flowpath as wall thickness is increased. In other words, 1 foot of 1/2" L copper has slightly less volume than 1 foot of 1/2 M copper.[citation needed]
We offer several financing options for new equipment, as well as our Priority Club Membership that grants members exclusive discounts, priority service, annual tune-ups, and much more. Plus, we make it our mission to give back to the community that's continually supported us for decades. Our team members regularly donate their time, money, and efforts to numerous charitable organizations, fundraising endeavors, and worthwhile causes throughout the Northern Virginia and suburban Maryland area.


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Postdoctoral Fellow in Lipid Biology and Metabolic Signaling in Skeletal Muscle Regeneration
- Employer
- University of Michigan Medical School
- Location
- Ann Arbor, Michigan
- Salary
- Salary will be based on experience, and adjusted up from the NIH postdoc pay scale.
- Posted
- April 16 2016
- Discipline
- Life Sciences, Biomedical Sciences, Cell Biology, Molecular Biology, Physiology, Stem Cells, Health Sciences, Regenerative Medicine, Translational Research
- Position Type
- Full Time
- Job Type
- Postdoc, Postdoc Fellowship
Position Description
A postdoctoral fellow position in skeletal muscle and connective tissue biology is available in the Mendias Lab (http://www.umich.edu/~cmendias). The lab is part of rich, interdisciplinary environment in the Department of Orthopaedic Surgery and Department of Molecular & Integrative Physiology at the University of Michigan Medical School. The postdoctoral fellow will participate in innovative projects looking at the contribution of lipid and metabolic signaling to muscle atrophy, fibrosis and inflammation using a combination of in vitro studies, genetically engineered rodents, mechanical loading studies and in biopsies from patients with chronic muscle injuries. The postdoctoral fellow will also have access to support and resources from the state of the art UM Metabolomics Core (http://mrc2.umich.edu). Work in our lab is currently supported by the NIH, Mark Cuban Foundation, Orthopaedic Research and Education Foundation, and Plastic Surgery Foundation.
Interested applicants should send their CV, a cover letter and a list of three potential references to Dr. Chris Mendias (firstname.lastname@example.com). Applications are also accepted through the UMjobs.org website, job opening ID 120232.
The position is expected to begin sometime in Summer 2016.
Required Qualifications
Must have a PhD and/or MD. Candidates should have a strong background in molecular and cell biology. Strong written and oral communications skills are necessary.
Desired Qualifications
Candidates with expertise in musculoskeletal biology or lipid biology are especially encouraged to apply.
Background Screening
The University of Michigan Medical School conducts background screening and pre-employment drug testing on job candidates upon acceptance of a contingent job offer and may use a third party administrator to conduct background screenings. Background screenings are performed in compliance with the Fair Credit Report Act.
U-M EEO/AA Statement
The University of Michigan is an equal opportunity/affirmative action employer.


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Translation from 'Comme des lions', Dominique Lormier
In our opinion, this must not be forgotten. This is why we want to introduce the French Army in one of the well-known modifications on WWII of Battlefield 2, Forgotten Hope 2.
(active, inactive and retired)
Special thanks to
Teams Contact ! and =UBF= for their support
Team 762
Team [F|H]
Team World at War
FH2 Community
No games, mods, engines, software or hardware appear to be published or in development by French Hope. We suggest you add games, mods, engines and hardware you are developing here (or update existing games, mods, engines and hardware on the site which should be linked with this Developer).
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Why has this not came out
Hi, this will come out soon, FH2 team and us are working on some maps atm, but i can't tell which one and when ;) but the progress is really good so the wait will be over soon !
Is there going to be a public event or something anytime soon? The mod is great and I want to play:)
Have you seen this:
Maybe if you could find the modeller, you might cooperate?
Nice stuff you got, but you really need some vehicles and stuff similiar. =)
Hey, do you know something about the french army weapons and equipment in the mod?
Superbe travail Bocquier, j'espère en voir encore de pareille carte
Vive la france, est la belgique :p
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Game of thrones: Fire and Blood Developer group...
Dedicated to bringing many 40K mods (ie. faction mods) to the Dawn of War 1 community.
A group of community-based Star Wars Empire at War: FoC MODS in Vkontakte
WeCreateStuff is a small community of geeks who work together on games, mods and all kinds of awesome stuff for the internet. We aim to keep on producing...
Call of Pripyat modding team. Currently the team consists of two designer and one coder, but you can apply for job. Currently developing Call of Chernobyl...
Group dedicated to improving and contributing to Dead Air.
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text: 65 Knyaz Boris I BlvdVarna, Bulgaria 9000View Large Map Phone: +359 52-989900Fax: +359 52-989902 Hotel E-mailHotel Website Classification 9 Deluxe Commission Rooms Rates 10% 55 $96-$224 Hoteliers! Is your information accurate? Update Here Overview A traditional hotel in Varna - Easy access to the Graffit Gallery, Regional Historical Museum, Varna Zoo and Dolphinarium. Details Year Built: 2011Check in Time: 2:00 PMCheck out Time: 12:00 PMNumber of Floors: 4 Graffit Gallery Hotel GDS Codes Amadeus GDS: OI VAR05ZGalileo/Apollo GDS: OI B0047Sabre GDS: OI 162685WorldSpan GDS: OI VARGG Graffit Gallery Hotel Rates & Policies Rate Policy: Daily in USDStandard Room: from $96-$152Suite: from $211-$224Credit Cards: Credit Cards Are AcceptedReservation Policy: Reservations must be guaranteed with a credit cardIncluded Meals: Continental Plan Graffit Gallery Hotel Room Amenities Amenities are in all rooms unless noted otherwise. Coffeemaker Rooms for Non-smokersInternet Access (High Speed & Wireless) Graffit Gallery Hotel Recreation Nearby Activities BeachGolf Course Graffit Gallery Hotel Hotel Services & Facilities Guest Services Laundry RoomLaundry/Dry Cleaning ServiceFree Onsite ParkingRoom ServiceWheelchair Access to Common/Public AreasCommon Area Internet Access (High Speed & Wireless)


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Collection of British Flag (25)
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Real estate '89.
As Hawaii approaches the 21st century, owning real estate in the Islands - whether a fee-simple home, a business, an investment in raw land or a commercial lease agreement - will become ever important. The growing population on Hawaii's limited land, coupled with continuing foreign investment, is making property ownership increasingly expensive.
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Developers working on real estate projects can benefit from expert consulting that answers a number of key question, including how to build for transportation, management and security considerations, and what clientele to target. Respected real estate firms with decades of experience in Hawaii can take the guesswork out of critical planning decisions.
On the following pages, some of Hawaii's leading professionals talk about themselves and the properties they represent.
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|special advertising section
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|May 1, 1989
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What are your chances of dating a celebrity
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The arrest for shoplifting that happened a decade ago will become news. The first months of were a steady drive in bliss and honey. We saw each other constantly and talked about the future. This time, our bond felt deeper. I was in love and overwhelmed. His team lost, but we still celebrated his favorite holiday in true Cinco de Mayo spirit: At a table filled with our friends, I stared at him. Not because he was wearing a silly oversize sombrero, but because I realized I was ready. I had finally arrived at a place where I could accept he was no longer an overgrown child. He was a grown-up, and he had made forgivable mistakes. We arrived back at his home, and I immediately, inexplicably, became violently ill all over his walls.
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What do you say? Um, wait, isn't he famous? Back off quietly and secretly hope they notice you're there with them 9 You and your date get pretty waisted at a bar.
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A celebrity may exist in a rarified bubble that skews their worldview.


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Original Research ARTICLE
Plant Density and Nitrogen Supply Affect the Grain-Filling Parameters of Maize Kernels Located in Different Ear Positions
- 1College of Agriculture, Nanjing Agricultural University, Nanjing, China
- 2State Key Laboratory of Crop Biology, College of Agriculture, Shandong Agricultural University, Tai'an, China
Although yield output of maize (Zea mays L.) has improved markedly over the last century, procedures for improving the grain-filling process remain elusive. Our aim in this study was to relate grain-filling variation in maize (including kernels in apical and middle positions in the ears) to plant density and nitrogen (N) application rate using a crossed experimental design. We also investigated changes in zeatin riboside (ZR), indole-3-acetic acid (IAA), abscisic acid (ABA), and gibberellic acid (GA) in the kernels during the grain-filling period. Two high-yield maize varieties cultivated extensively in China were field grown under normal (67,500 pl ha-1) and high (97,500 pl ha-1) densities, and supplied with low, normal and high (0, 180, and 360 kg N ha-1) concentrations of N. Kernel weight (KW), the maximum grain-filling rate (Gmax), the average grain-filling rate (Gave), and the kernel weight increment achieving Gmax (Wmax) were all significantly depressed under high density (HD) conditions, but increased N supply partially offset the losses. The apical kernels were more sensitive to density and N application rate than middle kernels. Correlation analysis indicated that plant density and N rate affected KW mainly by influencing the grain-filling rate. Variation in ZR, IAA, and ABA content tracked the variation in KW, but variation in GA content did not. Furthermore, the grain-filling parameters (closely related to TKW) had strong canonical correlation with the content of all hormones across the filling period and ZR content had the strongest relationship. Based on our study, high N supply is beneficial to optimize grain-filling parameters and improve KW of maize kernels under crowded condition.
A gradual increase in plant density has been a vital contributor to maize yield enhancement worldwide (Tokatlidis and Koutroubas, 2004; Antonietta et al., 2014; Testa et al., 2016). However, kernel number per individual plant (KNP) and kernel weight (KW) were decreased due to the exacerbated competition for light and nutrients among individuals under high densities (Tollenaar et al., 2006; Borrás et al., 2007; Boomsma et al., 2009; Al-Naggar et al., 2015). Maize yield is determined by the kernel number per unit area (KNA) and the kernel weight (KW) (Borrás and Otegui, 2001; Mansfield and Mumm, 2014). Once the number of kernels is established, the final KW becomes a main factor determining maize yield (Wang et al., 2012). Hence, improving KW under crowded conditions remains a challenging issue for further enhancements in maize production.
The final weight of maize kernels is generally determined by genotype characteristics (Reddy and Daynard, 1983), which varies greatly with the position on the ear (cob) (Xu et al., 2013), and is also affected by the growth environment (Gambin et al., 2006; Zhou et al., 2017). Sub-optimal conditions related to population density, nutrient status, temperature, drought, and light intensity may restrict kernel set and development (Borrás et al., 2003; Brekke et al., 2011), as well limit the maximum weight achievable (Ouattar et al., 1987; Borras and Gambin, 2010; Shi et al., 2013). For KW, maize grains are usually classified as inferior grains or superior grains, where the former are generally located on the upper part of the ear, and the latter are mostly located on the middle and lower parts of the ear (Chen Y.J. et al., 2013). KW varies with the rate of kernel growth and the duration of grain-filling (Gambin et al., 2007; Sadras and Egli, 2008), which are directly related to grain weight and yield (Borras et al., 2009; Chen T. et al., 2013). During much of the grain-filling period, grain dry weight increases at an essentially linear rate over time. This period of linear increase is known as the effective filling-period duration (EFPD) (Johnson and Tanner, 1972). Borrás and Otegui (2001) reported that KW was closely related to the rate of kernel growth during the EFPD rather than the duration of this period. However, Bolaños (1995) argued that improvements in maize yield were largely explained by the prolongation of the grain-filling period.
The grain-filling process can be accurately modeled with logistic equations, thereby generating a series of important grain-filling parameters, such as maximum grain-filling rate (Gmax), average grain-filling rate (Gave) and active grain-filling period (Xu et al., 2013). Besides, the whole grain-filling process could be further divided to three periods according to the shape of the logistic curve, i.e., (i) gradual increase period (GIP), (ii) rapid increase period (RIP), and (iii) slight increase period (SIP) (Wang et al., 2014). Previous work showed the period of maximal grain-filling rate was protracted as fertilizer dosage increased, and the whole process of grain-filling was prolonged (Cao et al., 2008). Xu et al. (2016) indicated the Gmax and Gave varied with different sowing dates, but the duration of the active grain-filling period did not. Variations of light intensity also affected grain yield by changing the Gmax (Shi et al., 2013).
Exogenous growth conditions and endogenous plant hormones are both key regulators of kernel development (Lv et al., 2017). Plant hormones play important roles in the development and enrichment of maize, wheat and rice endosperms (Yang et al., 2000; Zhang et al., 2009; Liang et al., 2017). For instance, high zeatin (Z) and zeatin riboside (ZR) content in the initial stage of grain-filling can promote kernel setting and enhance maximum endosperm cell division rate (Morris et al., 1993; Yang et al., 2002). Auxins, gibberellic acid (GA), and abscisic acid (ABA) also regulate grain development (Kende and Zeevaart, 1997; Hansen and Grossmann, 2000; Yang et al., 2006; Zhu et al., 2011). Xu et al. (2013) showed that kernel filling rate was strongly positively correlated with the concentration of Z+ZR, IAA, and ABA in the kernels, but significantly negatively correlated with concentration of GA3. Besides, the relative level of hormone in both superior and inferior florets were shown to determine the development of inferior florets (Wang et al., 2006). All early reports have shown that the grain-filling of kernels is controlled by hormones.
Improvements in grain-filling remain a substantial challenge for modern crop production systems with high yield output (Kato et al., 2007). Previous studies regarding grain-filling, especially the relationships between different pollination time and position, plant growth rate at different growth periods, various resource-sink ratio and kernel number set and KW have been established under diverse stress conditions (Zinselmeier et al., 1999; Andrade et al., 2002; Uribelarrea et al., 2002; Gambin et al., 2006). Nevertheless, many studies were concentrated on the early filling stage or the filling rate over the entire filling period. Besides, studies on grain-filling characteristics and KW were mainly focused on variety comparisons, ecological conditions and modifications of single cultivation practices. However, these previous studies showed no evidence on the response of grain-filling characteristics to contrasting stand densities and N availabilities. Information on the relationship between kernel ear position and grain-filling characteristics is also limited. Thus, we examined the grain-filling parameters of maize kernels (including apical and middle position kernels in the ears) under different plant densities and N supply rates using logistic modeling. Besides, the hormone content in grains were determined to analyze their relationships with grain-filling characteristics.
Materials and Methods
Trial Site and Conditions
Field evaluations were performed at the Shandong Agricultural University Experimental Farm in Shandong, China during 2014 growing season. This area has a semi-humid, warm temperate continental climate with monsoons. The average monthly maximum temperature, minimum temperature, total solar radiation, and precipitation during the experimental period are shown in Table 1. The soil at the site is neutral sandy loam, and the nutrient status of the top 30 cm before seeding consisted of 11.1 g kg-1 organic matter, 0.76 g kg-1 available N, 25.4 mg kg-1 available phosphate and 105.1 mg kg-1 available potassium.
Table 1. Monthly average maximum temperature (Tmax, °C) and minimum temperature (Tmin, °C) and monthly total solar radiation (SR, MJ m-2) and rainfall (mm) during the growing season (June to September).
Experimental Design
Two high-yield varieties planted widely in local production were selected as the testing materials, i.e., Denghai618 (521 × DH392) and Denghai605 (DH351 × DH382). The common density was 67,500 pl ha-1 and the N application rate ranged from 180 to 360 kg ha-1 in this location (Jin et al., 2012; Gao et al., 2017). Maize seeds were planted with hand planters on June 13 and harvested on September 29. Treatments consisted of two densities 67,500 pl ha-1 (low density, LD) and 97,500 pl ha-1 (high density, HD) and three N levels, 0 kg ha-1 (low N, N0), 180 kg ha-1 (normal N, N180), and 360 kg ha-1 (high N, N360). The experimental treatments were laid out in a split-split plot design, where the N rate was set as the main plot, and variety and density were set as subplot and sub-subplot with three replicates per treatment. The plot size was 12 m long × 3 m wide contained five rows and the space between rows was 60 cm. Plants within a row were placed 24.7 and 17.1 cm apart to achieve population density of 67,500 and 97,500 pl ha-1, respectively. Phosphorus (P2O5) and potassium (K2O) fertilizer were applied before sowing, at a rate of 90 and 120 kg ha-1, respectively. The N fertilizer was in the form of urea and applied as topdressing in two equal portions—one half at the jointing stage, and the other half at the large bell stage. Irrigation, weeds, diseases, and insect pests were controlled adequately during each growing season so that no factors other than density and N supply limiting growth.
In each plot, at least 50 representative plants with robust and uniform growth were labeled and bagged before silking. After the silks were fully exerted, artificial pollination was conducted uniformly on the same day. The shoot bags were replaced after pollination and removed only after pollen stopped shedding. All data were obtained from three rows of plants in the center of the cultivation plots.
Sampling and Measurements
Grain-Filling Traits
Five labeled ears were sampled from each plot to determine grain-filling parameters at 10, 20, 30, 40, and 50 days after pollination (DAP). Kernels from the apical 5–25% files were collected and mixed to provide samples of apical kernels (inferior kernels). Kernels from the 40 to 60% files (counted from the apex to bottom) were collected and mixed to provide middle position grain (superior kernel) samples. We sampled 200 kernels from the five ears and randomly divided them into two equal portions. One portion was frozen in liquid N for 10 min and then stored at -80°C. The second portion was held in an oven for 30 min at 105°C and then dried to constant weight at 80°C.
Grain-filling process was fitted by the logistic equation W = A/ (1+Be-Ct). Where W refers to the kernel weight, t refers to days after pollination (the day of pollination set as t0 = 0), A refers to ultimate growth mass, B refers to primary parameter and C refers to growth rate parameter, respectively. The grain-filling parameters of the tested maize varieties were calculated as follows.
Occurrence time of maximal grain-filling rate (Tmax) = lnB/C;
Kernel weight increment achieving maximum grain-filling rate (Wmax) = A/2;
Maximum grain-filling rate (Gmax) = (C × Wmax) [1-(Wmax/A)];
Initial filling power (R0) = C;
Active grain-filling period (P) = 6/C.
The whole process of grain-filling was divided into three periods according to the shape of the logistic curve: (i) gradual increase period, (ii) rapid increase period, and (iii) slight increase period. The first and second order derivatives of the logistic equation were calculated to determine the filling parameters of these three grain-filling periods. The start time of peak grain-filling (t1) = (ln B-1.317)/C, the grain weight w1 = A/(1+Be-Ct1); the end time of peak grain-filling (t2) = (ln B+1.317)/C, the grain weight w2 = A/(1+Be-Ct2); the effective grain-filling period (when the grain weight reached 99% A, t3) = (ln B+4.59512)/C, the grain weight was w3. The grain-filling parameters of the three grain-filling period were calculated as the following equations.
The gradual increase period (T1) = t1-t0;
The rapid increase period (T2) = t2-t1;
The slight increase period (T3) = t3-t2;
The increment in grain weight during gradual increase period (W1) = w1-w0;
The increment in grain weight during rapid increase period (W2) = w2-w1;
The increment in grain weight during slight increase period (W3) = w3-w2;
The average grain-filling rate during gradual increase period (G1) = W1/T1;
The average grain-filling rate during rapid increase period (G2) = W2/T2;
The average grain-filling rate during slight increase period (G3) = W3/T3.
Besides, the average grain-filling rate during the whole grain-filling period was calculated as:
Average grain-filling rate (Gave) = W3/t3.
Hormone Content
The endogenous ABA, IAA, GA, and ZR were extracted with 80% (v/v) methanol and quantified by enzyme-linked immunosorbent assay (ELISA) according to previous studies (Liang et al., 2017). The ELISA kits were provided by the Phytohormones Research Institute, China Agricultural University. The recovery rates for ABA, IAA, GA, and ZR were (90.2 ± 4.1), (86.1 ± 4.5), (79.4 ± 5.3), and (93.4 ± 6.5)%, respectively.
Grain Yield, Kernel Weight, and Kernel Number
At physiological maturity, 30 ears from the center three rows of each plot were harvested and air-dried to investigate yield, KNP and 1000-kernel weight (TKW). Grain yield (Mg ha-1) was expressed at 15.5% moisture.
Statistical Analysis
The Curve Expert 1.3 software was used to fit the grain-filling process equation and obtain parameters A, B, and C. Analysis of variance (ANOVA) was performed with SPSS ver. 18.0 (SPSS Institute, Inc.). Duncan's multiple range test was used to evaluate differences among treatments, and the significance level was set at the 0.05 probability level. Pearson correlation analysis and canonical correlation analysis were also performed. Figures in the article were plotted using SigmaPlot ver. 12.0 and GraphPad Prism ver. 7.
Grain Yield and Yield Components
Density, N rate, maize variety and the interactions between them (the density × variety interaction term was not included) had significant effects on GY (Table 2). The highest GY for variety DH618 was obtained under a combination of high density (HD) and high N rate (360 kg N ha-1, N360). The highest yield of variety DH605 was obtained with a combination of HD and a N rate of 180 kg ha-1 (N180). The GY response of DH618 to N (N180, N360, vs. N0) was greater under HD (GY increases of 38.1 and 46.3% with N180 and N360 fertilization, respectively) than at low density (LD) (GY increases of 31.5 and 32.4%, respectively). In DH605, the respective GY increases under HD were smaller: 22.2% (N180) and 22.5% (N360).
Table 2. Effects of density and N rate on grain yield (155 g kg-1 water content), kernel number and kernel weight.
Density and N rate had significant effects on KNP and 1000-kernel weight (TKW), but the density × N interaction term was not significant (P > 0.05). Increases in stand density were accompanied by decreases in KNP and TKW, but the values of both of these response parameters increased with increasing N rate, especially under HD (KNP values for DH618 increased on average by 27.9 and 17.9% under HD and LD, respectively). The KNP response of DH618 (average 23.0%) exceeded that of DH605 (average 9.9%). The difference in TKW between the two densities as the N rate increased was less marked than the KNP response. The TKW responses of DH618 exceeded those of DH605. For example, in DH618, the TKW value in the N360 × HD treatment increased by 7.9% in comparison with the N0 treatment, whereas the TKW proportional response to the same treatment was 4.8% in DH605.
Grain Weight Dynamic
Density, N rate and variety significantly affected KW in both the apical and middle ear positions (data not shown). The density × N rate interaction term was significant after 10 DAP for middle position kernels, but not for the apical grains. The KW of middle position kernels was higher than that of apical kernels over the whole grain-filling period. In each of the grain-filling periods, increased density reduced KW, while increased N application rates improved KW values (Figure 1), as expected. The responses of kernels in different ear positions were always different. Apical kernels responded more strongly to density and N supply than middle position kernels. Thus, in variety DH618, the KW values of apical and middle position kernels under HD were reduced on average by 8.6 and 5.9%, respectively, in comparison with the LD treatment. The respective average KW values in the high N treatment (N360) were 7.7 and 6.3% above the values in the N0 treatment. The response to N (N180, N360, vs. N0) was greater at HD. For example, under HD, the KW values of middle position kernels of DH618 at 30 DAP were 5.3% (N180) and 7.0% (N360) higher than those in the N0 treatment, whereas the respective increases under LD were 3.4% (N180) and 4.6% (N360).
Figure 1. Effects of density and nitrogen application rate on the grain dry weight and grain-filling rate of (A,B,E,F) upper kernels and (C,D,G,H) middle kernels (A–D) in Denghai618 and (E–H) Denghai605 during the 2014 growing season. LD and HD refer to low density and high density, respectively. N0, N180, and N360 represent nitrogen rates of 0, 180, and 360 kg ha-1, respectively.
Grain-Filling Characteristic Parameters
We divided the grain-filling parameter values under different D × N combinations by the value for the LDN0 treatment. These ratios are listed in Supplementary Table S1 and are presented in Figure 2. At 10 DAP, the different density and N rate combinations showed no significant effects. However, after 20 DAP, the differences in grain-filling rates among treatments became obvious. Progress in grain-filling was well-simulated by the Richards model: all correlation coefficients were > 0.99 (Table 3). The maximum grain-filling rate (Gmax), the kernel weight increment achieving Gmax (Wmax), average grain-filling rate (Gave) and the active grain-filling period (P) all decreased with increased density, but increased with the N supply rate. The initial grain-filling potential (R0) and the day on which the maximum grain-filling rate (Tmax) reached responded differently. Compared with the values for superior kernels, the Gmax, Gave, and P-values of inferior kernels were depressed. However, apical kernels responded more strongly to N supply. Gmax values in treatment N360 (DH618) were elevated by 8.6 and 5.6% in apical and middle position kernels, respectively, above the values in the N0 treatment. The grain-filling rate was higher in DH618 than in DH605, but the active grain-filling period of DH618 was shorter.
Figure 2. The variations of grain-filling characteristic parameters under different density and N application rate combinations. The results are the ratios of grain-filling characteristic parameters values under different D × N combinations to LDN0 for each variety (i.e., ratios of LDN180/LDN0, HDN180/LDN0, et al.), where LD and HD refer to low density and high density, respectively. N0, N180, and N360 represent nitrogen rates of 0, 180, and 360 kg ha-1, respectively. Gave, average grain-filling rate; Tmax, the day reaching the maximum grain-filling rate; Wmax, kernel weight increment achieving maximum grain-filling rate; Gmax, maximum grain-filling rate; R0, initial grain-filling potential; P, active grain-filling period; T1, grain-filling duration of gradual increase period; W1, increased grain weight of gradual increase period; G1, mean grain-filling rate of gradual increase period; T2, grain-filling duration of rapid increase period; W2, increased grain weight of rapid increase period; G2, mean grain-filling rate of rapid increase period; T3, grain-filling duration of slight increase period; W3, increased grain weight of slight increase period; G3, mean grain-filling rate of slight increase period.
Table 3. Effects of density and N rate on grain-filling characteristics parameters of maize kernels at upper and middle position.
Density increases prolonged the duration of the GIP in most cases, but shortened the durations of two other periods on the logistic curve. The GIP, RIP and SIP durations were all prolonged in both inferior and superior kernels of the two varieties by increasing the N supply rate (Table 4). The increases in KW of inferior kernels through all the three filling periods were smaller compared with the increases in superior kernels. The duration of the apical kernel RIP and SIP were shorter, but the duration of the GIP was longer in comparison with middle position kernels. In addition, the duration of the DH618 GIP was shorter than the DH605 GIP, but the grain-filling rate and the increases in grain weight during the GIP were higher in DH618.
Table 4. Effects of density and N rate on characteristics parameters of the three grain-filling phases of maize kernels at upper and middle position.
Variations in Hormone Content
During the grain-filling process, the overall trends in IAA, ZR, and ABA content in the grains showed unimodal curves. The GA content declined continually in all treatments (Figure 3). Plant density and N rate both had significant effects on hormonal content during the whole filling period. The density × N rate interaction was significant for most hormone content (Supplementary Table S2). Density and N rate did not alter the general trends of any of the hormones over time. At each grain-filling stage, IAA and ZR content were higher in middle position kernels than in the apical kernels, but the GA content was higher in the apical kernels. Before 30 DAP, the concentration of ABA was much lower in the apical kernels than in the middle position grains. The hormone content in apical kernels responded more strongly to density and N supply than in middle position kernels. For example, the Z+ZR content at 20 DAP was reduced by average 8.8 and 3.7% (HD vs. LD), but were increased by average 17.3 and 5.9% (N360 vs. N0) for inferior and superior kernels, respectively. The response to N (N180, N360, vs. N0) was also elevated in the HD treatment. Thus, the Z+ZR content at 20 DAP (middle position kernels of DH618) was increased by 2.8% (N180) and 3.5% (N360) under LD, and by 6.2% (N180) and 7.7% (N360) under HD.
Figure 3. Effects of density and nitrogen application rate on the content of zeatin riboside (ZR), indole-3-acetic acid (IAA), abscisic acid (ABA), and gibberellic acid (GA) in (A,C,E,G,I,K,M,O) upper kernels and (B,D,F,H,J,L,N,P) middle kernels of the two varieties during 2014 growing season. LD and HD refer to low density and high density, respectively. N0, N180, and N360 represent nitrogen rates of 0, 180, and 360 kg ha-1, respectively.
Correlation Between 1000-Grain Weight and Grain-Filling Parameters
We examined the relationships between grain-filling parameters in different ear positions and the TKW (Table 5). Correlation analysis showed that Gave, Wmax, Gmax, the grain-filling rate and KW increased during the GIP, RIP, and SIP were significantly positively correlated with TKW at physiological maturity in kernels at both ear positions. We also found a significant negative relationship between KW and the duration of the GIP in middle position kernels.
Table 5. Pearson correlation of 1000-kernel weight (TKW) and filling parameters of maize kernels at different ear positions.
Correlation Between Grain-Filling Parameters and Hormone Content
Canonical correlation analysis was used to examine the relationships between hormone content (X) and grain-filling parameters (Y). Hormone content at 10, 20, 30, 40, and 50 DAP were set as variables x1, x2, x3, x4, and x5, respectively. Gave, Wmax, Gmax, G1, G2, and G3 were set as y1, y2, y3, y4, y5, and y6, respectively. As shown in Table 6, the first canonical correlation coefficients were all statistically significant, indicating that the correlations between the grain-filling parameters and hormone content can be described by the first pair of typical variables (except the ABA content of superior kernels). The linear combinations of the first pair of canonical correlation models are shown in Table 7. From the coefficients for each variable, we were able to show that the grain-filling parameters of apical kernels were mainly determined by the content of ZR at 40 DAP, IAA at 20 DAP, ABA at 10 DAP and GA at 40 DAP, whereas content of ZR at 50 DAP, IAA and ABA at 20 DAP, and GA at 10 DAP were the main factors determining the grain-filling parameters of the middle position kernels. Most information of X and Y were concentrated in the first pair of canonical variables (L1 and M1, Table 8). Among the hormones examined in our study, ZR had the strongest correlation with grain-filling parameters of both inferior and superior kernels (86.0 and 79.6% of the ZR variation may affect variation in grain-filling parameters in inferior and superior kernels, respectively; 90.2 and 91.9% of the variation in grain-filling parameters were related to ZR content changes in the inferior and superior kernels, respectively).
Table 6. Canonical correlation coefficient between grain filling parameters and hormone content and the significance testing.
Table 8. The proportion of variable variation can be explained by canonical variable (redundancy analysis).
Although maize GY is determined mainly by KNA, variations in KW are also influential (Zhou et al., 2016). Grain-filling is a critical physiological process that determines the KW and yield of corn kernels. Poor grain-filling limits the potential GY (Yang and Zhang, 2010) and is among the major scientific and practical issues that should be addressed to improve agricultural production. Appropriate plant densities and N fertilization rates are important elements of efficient management that have contributed to maize yield improvement (Wei et al., 2017). We showed in this study that density, N rate and their interaction affected KW during the whole filling period, while the impact of density tended to be stronger. The variation in KW between different floret positions of the maize ear was substantial and mostly related to changes in the grain-filling rate. Furthermore, the low KW values in apical kernels were exacerbated under HD, though the yield potential under HD was nevertheless 9.8% higher than that under LD. Taken together with the strong response of apical kernels to N rate (especially under HD), we concluded that apical grains were more sensitive to environmental factors than middle grains. Hence, promoting grain-filling of grains at apical position will be key for improving the KW of maize.
The KW is determined by the grain-filling rate, the effective filling duration, or both (Zhou et al., 2017). In our study, Gave, Gmax and the filling rate during GIP, RIP, and SIP were all positively correlated with KW, but we found no obvious correlation between KW and the active grain-filling period in neither superior kernels nor inferior kernels (Table 6). Thus, changes in the filling rate were mainly a result of variation in KW under different density and N rate combinations. Li et al. (2000) showed that difference in KW within the same maize variety was determined by the grain-filling rate, whereas KW variation among varieties was determined by grain-filling duration. We attributed KW differences between the two maize varieties that we tested to differences in filling rate, rather than filling duration. Unlike apical kernels, the duration of the GIP in middle position kernels was strongly negatively correlated to KW. A mechanism for shortening the duration of the GIP whilst maintaining the filling rate could prolong the duration of the subsequent RIP relative to the SIP. This procedure could increase KW.
The processes of cell division, differentiation, enlargement, and enrichment of inclusions during the development of maize kernels are controlled by a range of hormones operating during different filling periods (Yang et al., 2001; Artlip et al., 2010). In the current study, density and N level did not change the regulation period of the hormones in grains, however, hormone content varied significantly through kernel development. HD decreased the highest values, and accelerated reductions in ZR and IAA content, whereas increased N application offset these declines, especially under HD. The strong canonical correlation between grain-filling parameters and ZR content indicated that density and N rate combinations affected grain-filling progress largely by adjusting the content of ZR, which regulated the rates of endosperm cell division and grain filling (Ahmad et al., 2018). Additionally, we also showed that the ZR and IAA content of apical kernels were significantly lower than the content in middle position kernels. Hence, we attributed the lower KW in the apical kernels to their lower ZR and IAA content, these low hormone levels constrained the proliferation of endosperm cells (Wang et al., 2006), reduced the sink capacity, and ultimately decreased KW. ZR and IAA are considered to be the main regulators of endosperm cell development, whereas ABA likely has a vital regulatory role in the process of photosynthate transport and accumulation in the grain (Hansen and Grossmann, 2000; Travaglia et al., 2007; Qin et al., 2013). We found that the ABA concentration in the kernels was reduced by plant crowding, but enhanced by increased N availability, in agreement with the findings of Gawronska et al. (2003). Furthermore, we determined that about 75% of the variation in grain-filling parameters was related to variation in ABA content in both apical and middle position kernels. The ABA content during the early filling period (10–20 DAP) was most strongly positively correlated with the grain-filling parameters, as shown in previous studies on maize, wheat, and rice (Xu et al., 2007, 2013; Zhang et al., 2009; Lv et al., 2017). The combination of elevated ABA concentration after 30 DAP and the low KW in the HD × N0 treatment also suggested that maintaining a high level of ABA during the late grain-filling stage may negatively impact the progress of grain-filling.
Unlike the other hormones, the GA levels in both apical and middle position kernels were highest at 10 DAP, when the embryo was enlarging rapidly. In constract to the ZR content, GA content increased when density was increased, but declined with increased N application. The concentration of GA in the apical kernels exceeded that in the middle position kernels. Earlier studies have indicated that GA in the kernels played an important role in the rapid amplification of endosperm cells. However, we found that the GA content in the HD × N0 was maintained at a high level over the whole filling period, suggesting that elevated GA content adversely affected the improvement of KW. This outcome is congruent with earlier work (Xu et al., 2013), showing that GA can enhance the activities of α-amylase and other hydrolytic enzyme that promote the decomposition of starch (Yang et al., 2001). Thus, changes in ZR, IAA, ABA, and GA content in grains were all significantly correlated with the grain-filling parameters. Further work is needed to explore the mechanisms controlling (i) hormone content variation under different density × N rate combinations, and (ii) hormone distribution within maize ears over the whole grain-filling period.
In short, variation in grain-filling rate was the main factor affecting TKW under different density × N rate combinations. The apical kernels were more sensitive to density and N application rate than middle position kernels. High N input weakened the reduction in KW, Gmax, Gave, Wmax, RIP, and SIP durations under high density. The grain-filling parameters (closely related to TKW) had strong canonical correlations with the content of all hormones across the filling period, and ZR content had the strongest relationship. Therefore, we conclude from our results that high N supply is beneficial to optimize grain-filling parameters and improve KW of maize kernels under crowded condition.
Author Contributions
SW designed the work, carried out all experiments and data analysis and drafted the manuscript. XW performed field experiments and sampling and helped drafting the work. SD conceived the study and planned the experiments. GL and YQ helped draft the manuscript. DJ helped draft and revise the manuscript. All authors read and approved the final manuscript.
This research was funded by the Natural Science Foundation of Jiangsu Province (Grant No. BK20170720), the China Postdoctoral Science Foundation funded project (Grant No. 2017M611832), National Natural Science Foundation of China (Grant No. 3117 1497), the National Basic Research Program of China (973 Program, Grant No. 2011CB100105), the National Food Science and Technology of High-yield Program of China (Grant No. 2011BAD16B09), the Special Fund for Agro-scientific Research in the Public Interest of China (Grant No. 20120306), and supported by the 111 Project (Grant No. B16026).
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.
The authors gratefully acknowledge Liyuan Hou, Meifeng Zeng, and Ting Liang for helping us to perform the study. They would also like to thank Mr. Mingcai Xu and all the workers in their farm for managing the field experiments.
Supplementary Material
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fpls.2019.00180/full#supplementary-material
TABLE S1 | The data used in Figure 2.
TABLE S2 | Analysis of variance of hormone contents in maize grains under different density and N rate conditions.
G1, mean grain-filling rate of gradual increase period; G2, mean grain-filling rate of rapid increase period; G3, mean grain-filling rate of slight increase period; Gave, average grain-filling rate; Gmax, maximum grain-filling rate; KW, kernel weight; P, active grain-filling period; R0, initial grain-filling potential; T1, grain-filling duration of gradual increase period; T2, grain-filling duration of rapid increase period; T3, grain-filling duration of slight increase period; TKW, 1000-kernel weight; Tmax, the day reaching the maximum grain-filling rate; W1, increased grain weight of gradual increase period; W2, increased grain weight of rapid increase period; W3, increased grain weight of slight increase period; Wmax, kernel weight increment achieving maximum grain-filling rate.
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Keywords: canonical correlation analysis, grain-filling characteristic, kernel position, hormone, logistic curve, maize
Citation: Wei S, Wang X, Li G, Qin Y, Jiang D and Dong S (2019) Plant Density and Nitrogen Supply Affect the Grain-Filling Parameters of Maize Kernels Located in Different Ear Positions. Front. Plant Sci. 10:180. doi: 10.3389/fpls.2019.00180
Received: 18 June 2018; Accepted: 05 February 2019;
Published: 01 March 2019.
Edited by:Abraham J. Escobar-Gutiérrez, Institut National de la Recherche Agronomique (INRA), France
Reviewed by:Axel Tiessen, Centro de Investigación y de Estudios Avanzados (CINVESTAV), Mexico
Michael Gerard Muszynski, University of Hawai'i at Mānoa, United States
Copyright © 2019 Wei, Wang, Li, Qin, Jiang and Dong. 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) and the copyright owner(s) 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: Dong Jiang, firstname.lastname@example.com Shuting Dong, firstname.lastname@example.com


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