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Finding macro wildlife
|A spider on the hunt is one of countless scenes available to photograph, if you know how to find them.
The arthropod world is virtually endless. With over a million recorded species - believed to be but a fraction of the actual total - you can shoot ten different species every single day of your life and never get close to documenting them all. Finding them in their native habitat exhibiting natural behavior, however, can be a challenge for those new to wildlife macro photography. In this article I'll walk you through the task of locating and handling macro subjects in the field.
|Elad and Bruno admiring two spiders in their natural habitat. The key for any successful macro photographer is to be able to closely observe a subject without disturbing its natural behavior.
First, however, I should point out that insect activity is influenced by climate and seasonal conditions, which of course vary greatly around the globe. Here I'll speak to my experiences photographing in countries with 'normal' winter/summer climates in which mornings are chilly and midday is considerably warmer.
The first order of business is getting to know the best seasons of the year to find and photograph insects. While there are insects present year round in most climates, it is in springtime that the populations truly boom. Spring, in most species, is the time of year when the adult hatches from its cocoon, and starts frantically looking for food and for a mate. The fields are absolutely packed with invertebrates of all species, flying, hunting, mating - doing everything we want them to do to create an interesting shot.
|Robber fly mating is precarious. The male often waits for the female to catch prey before he approaches, so he doesn't become her next meal himself!
During summer months, insect numbers dwindle noticeably but there are still lots of subjects available. In the fall and winter, however, things are usually not as good. Most adult insects will have died out and the newborns will be in one of their larval stages.
One of the most common questions I get is, 'Where are good places for finding little critters?' The answer is simple. Insects and other invertebrates are everywhere. They exist in almost any environment. And the more remote and wild the location, the higher the quantity and diversity.
The best locations though, are usually ones with damp earth, vegetation, and most importantly – water. I always recommend exploring areas near lakes or in fields with low growth. A body of water with a field beside it is just perfect. Some insects (such as dragonflies) are only found close to water, since their larval stage is fully aquatic.
|A butterfly found in low vegetation.
|Be careful when you walk in the field: there are countless meticulously-built spider webs!
Many species of course, have no problem with a drier environment; robber flies and mantises, to name just two. Fields of low growth vegetation are much easier to maneuver in and generally allow for more pleasing compositions. All else being equal, you'll likely have much more success in a sparse poppy field than a densely packed sunflower plantation for example.
Time of day
Arguably the most important consideration of all is knowing what times of the day are most conducive to locating and shooting invertebrates. Time of day has a huge impact not only on subject activity, but obviously on the quality of light as well.
The best time to shoot non-active arthropods is very early in the morning. Why? Because insects are less active when the temperature is lower. Under crisp early morning conditions, not only will insects hold 'poses', they can be easily manipulated and repositioned by hand without causing them any harm.
Natural behavior
Finding insects is one thing. Getting close enough to photograph them at macro-required distances while they're exhibiting interesting behavior is another skill altogether. But its not as difficult as it may seem if you know some basics.
The first thing to know is that insects are highly motion-sensitive. Making sharp, abrupt movements close to an active insect will likely cause it to flee. The rule of thumb here is to plan your movements carefully and perform them as if in slow motion. In addition, you should know that insects' eyes are ultra-sensitive to changes in light. This is clearly a survival response, since a sudden shift from light to dark could indicate a predator is approaching. By taking care to move toward the subject without casting a shadow near it, you increase the chance of it staying put for the shot.
|An active butterfly can still be approached if we make sure we don't frighten it by casting a shadow or making sudden movements.
Finally, it's important to understand the types of activities in which insects are more prone to remain stationary. A robber fly, for example is less likely to fly away if it's carrying prey. A female spider will not run away from her egg sack, even at the expense of her life. Knowing even a few little details like these enables you to get very close to your subject.
|A robber fly carrying prey is easy to get close to, if you approach it slowly.
|A female crab spider guarding its egg sack will never leave it, allowing an extreme magnification shot with minimal effort.
With these tips in mind, you can greatly increase your chances of creating unique and impressive wildlife macro imagery.
For more information on macro photography take a look at Erez's other articles:
The what and why of wildlife macro photography
What we want in a macro shot - Detail
What we want in a macro shot - Background
What we want in a macro shot - POV and special scenes
Macro photography: Understanding magnification
Depth of Field in Macro Photography
Composition Basics in Macro Photography
Erez Marom is a professional nature photographer and photography instructor based in Israel.
In January 2013, Erez will lead a winter landscape photography workshop in Iceland, where you can experience and shoot incredible icy sceneries. See Erez' Iceland gallery here. You can see more of Erez' work at Erezmarom.com and follow him on his Facebook page and deviantArt gallery.
Apr 8, 2015
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Jan 14, 2015
The tip about shooting the insects in the morning works like a charm
very nice picture
good pitures i like the nature and pros of the pictures
Would love an article about lighting in this series.
Gorgeous pics man. Never thought insects could be as beautiful as these. They are like some of the most beautiful birds around. I am not sure whether it is the quality of the cameraman or the modern cameras that makes such beautiful pics. But they are awesome nevertheless.
Nice article.
I can confirm that the ONLY (semi) successful dragonfly shoot I ever did (it's really not my thing, but the opportunity presented itself), was frighteningly early in the day:
Roel Hendrickx
Like your advise OldArrow.
Nicely presented set of good advices! Let me add an observation.
It is of utmost importance to emit peaceful, soothing and admiring thoughts when approaching any animal. Sometimes it makes all the difference! If you think fear, hunt, catch or similar thoughts, animals are very likely to sense those, and will probably try to mantain safety distances.
I learned this underwater. If you enter the water with a speargun, the sea in front of you will empty itself - not that the fish know about the harpoon, but they can sense "hunter's intentions". On the other hand, when you're carrying a camera, your intentions will be sensed as harmless, and the fish will not be alarmed at all.
I can't say that I understand the mechanisms involved, I only know it works that way, every time. There are several theories about that, but they are only theories. We may never know how... but the main thing is, it works.
Good article, well done
one thing that I have learned from my own experience is that most insects return to where after a while, so if you stay near the point where you saw an insect, there is a fair chance that it will come back after some time
great point. when i shoot dragon fly's, i find a place where there are a ton of them. i plop down in the middle and slowly wait for them to return . after a while they forget i am there. what doesn't work is chasing them around. find where they want to be, settle in and let the bugs come to you.
I used to take classes in entomology, so I've picked up a few pointers on finding insects that I thought would be worth sharing here.
1: Life prefers edges. You'll find a much greater variety of insects at the border between a forest and a field than you will in the middle of either. Treelines, pathways, and edges of streams are all excellent places.
2: Dress for the occasion. Wear bright colors and some fragrance, and the bees and butterflies are likely to come to you.
3: If you're looking for the bugs themselves, you'll have a rough time of it. Look instead for evidence of bugs. Chewed leaves, withered plants, dark spots on trees, rolled up leaves; most anything out of place.
4: Patience is key. Sometimes it's best to just sit by some flowers and wait for the bugs to come to you. Try not to panic when a few inevitably use you as a place to rest. Use slow, deliberate movements.
5: For nocturnal insects, light traps will make things really easy.
Good luck, Everyone!
These are good comments, thanks. I would add that I think bugs will become more easily observable over time.
I haven't ever hunted bugs. However, other times I have been in areas to observe animals and such I have not at first seen much. Over time, it became easier for me to see different animals.
Yes, useful, but we're still waiting for the big secret of the series - how do you get them to stay still while you focus stack?
try to find sleeping insects or as mentioned above guarding spiders. I like eating insects to.
for more closeups visit blog.mdsign.nl
its eyes are closed! duh!! ;-)
I always use focus stacking technique. The secret is you should get up before the sun come up. I often got up at 4 o' clock.
Some picture if you are interested:
What about hypnosis? No,really,the bussiness about wearing bright colours it trully works. I wore a long sleeve tiny flowers design shirt at a park in the Amazon,and, guess what, people arond me had a ball of fun taking pictures of a butterfly that would not stop crowling all over me. It wanted to make sure every flower in the shirt was scrutinized. Seriously!
Thank you Patrik, i look at your very nice gallery, but i liked the photos of you in the field, sure here a photo is more explanatory than thousand words about how to go with the setting. Good work, boy and thank s for sharing.
Good tips and wonderful images Erez. Your web site is very well done too. The gallery pages are very inspiring. Thank you for sharing both knowledge and images.
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A roll of film discovered in an Argus C2 from an Oregon Goodwill turned out to contain some incredible images – and has been re-united with the original owner's family.
Nikon's 28mm F1.4E ED appears to roundly complete the company's updated lineup of fast, professional prime lenses. We've already seen some initial images from a Nikon ambassador, but we've worked through a gallery of our own, with a lens of our own over the past week. Take a look.
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Post-Matric Scholarship Scheme
All India Scholarship Scheme
Steps to fill Scholarship Form Online
Eligibility Criteria for Post-Matric Scholarship Scheme
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subset: commoncrawl


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Cats get injuries that need to be treated with ointments just like humans. However, before applying an ointment, you should always take your cat to the vet to get it checked out for any internal injuries. Generally, vets will prescribe an ointment, but there are some over-the-counter ointments you can buy from pet stores to fix cuts and bug bites. Be prepared to deal with a struggling cat, because most do not like any ointment application.
Things You'll Need
- Gauze
- Antibacterial soap
- Scissors
- Antibacterial ointment
- Bandage
- Cardboard
- String
- Collar
- Hydrogen peroxide
- Betadine
- Treats
- Helper (optional)
Wrap the cat up in a towel to prevent it from wiggling away or scratching. Locate the injury on its body. Apply pressure to the injury with a gauze. Wait for the wound to stop bleeding before taking off pressure.
Clean your hands with antibacterial soap. Also, clean a pair of scissors with the soap. Rinse the scissors off with some hydrogen peroxide. Cut the fur around the wound carefully. You may need to have someone hold the cat when you trim the fur.
Wash the wound off with lukewarm tap water. Inspect the wound. If the wound is from an animal bite or is over an inch long, take your cat to the animal hospital.
Dab antibacterial ointment, such as Neomycinon or Triple Antibiotic Ointment, on the wound. Wrap the wound with a sterile bandage that sticks to itself. Using tape on a cat will cause its hair to stick to the bandage and will not be comfortable for the pet.
Make a protective collar for your cat so it does not try and bite or lick the wound. Cut a circle, 12-inches in diameter, out of a cardboard box. Make a hole in the center of the cardboard circle that is four to five inches in diameter--basically, the size of your cat's neck. This will become the cat's protective collar. Punch two small holes at the bottom of the cardboard collar and tie a piece of string to each hole. Slip the cardboard collar over the cat's head. Tie the strings attached to the cardboard collar onto the cat's existing collar. Remove the cardboard collar when the cat wants to drink or eat.
Clean the wound after two days with a mixture of three percent hydrogen peroxide and five parts water. Only clean the skin once with the mixture, because it can damage skin tissue and impede healing. Cleanse the wound with a mixture of one part Betadine and 10 parts water after that until it heals. Apply ointment daily.
Tips & Warnings
- Keep your cat occupied while tending to its wound by feeding it its favorite treats.
- Monitor your cat for a fever, because it is a sign of infection.
Related Searches
- Photo Credit Comstock/Comstock/Getty Images
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Preservation New Jersey • 1867 Sanctuary at Ewing • PO Box 7815• West Trenton, NJ 08628
Physical (GPS) address: 101 Scotch Road, Ewing, NJ 08628 Map and Directions
609-392-6409 firstname.lastname@example.com
Grand Opening Festival
Contemporary Music Concert Series:
Autumn Cabaret
Catrina Faith Orsi
Andrew W. Monath
Friday, October 21, 2016, 8:00 pm
at the 1867 Sanctuary at Ewing
Preservation New Jersey is pleased to present Catrina Faith Orsi and Andrew W. Monath in celebrating life in a Fall Cabaret in the acoustically beautiful and historic landmark 1867 Sanctuary at Ewing!
Catrina and Andrew are fun, energetic, youthful and have stories to tell through their music, Catrina as a young mom and Andrew as a performer and teacher.
Songs will include such favorites as: Man of La Mancha, Crazy, Fallin', Laughing Matters, Wizard and I, There's a Fine, Fine Line, Hopelessly Devoted to You, You Can Always Count on Me, Romancing the Blues, Big Time, and We Live on Borrowed Time.
ANDREW W. MONATH earned a bachelor's degree in Music Education from Westminster Choir College and a Master's Degree in Integrated Arts Education from Arcadia University with a concentration in Musical Theater. He taught public school music for eighteen years, spending three years as the Choral and Drama/Musical Director at Lyndhurst High School in Bergen County, NJ. His previous teaching positions have included Choral Director and Drama Director at Highland Park Middle and High School and 12 years as a Middle School Choral Director in Cheltenham School District where he built the middle school choral program to over 300 voices and established a nationally renowned musical theater program for middle school students. His choirs have sung for festivals and other community events as well as the National Anthem at Citizen's Bank Park for a Phillies Game. He has been a professional Church Musician for over 25 years. He is beginning his second year as the Minister of Music at The Reformed Church of the Ascension, United Church of Christ in Jeffersonville, PA. Prior to the he served as Organist and Music Director at Arch Street United Methodist Church in Philadelphia and before that he served as Organist/Music Director at Asbury United Methodist Church in Norristown, PA for 12 years. At Asbury he has helped the Chancel Choir increase in numbers as well as established the Sanctuary Concert Series, formed the Youth Choir and a Community Outreach Choir.
In addition to his work in Choral Music, he has directed and produced over fifty plays and musicals with middle school, high school and college aged students. Mr. Monath has been an adjunct professor at Arcadia University where he taught "Acting Styles in Musical Theater" and served as Musical Director in the Theater Department. For seven years he served as the Assistant Conductor for VOICES Chorale in Princeton, NJ. Mr. Monath is a member of ACDA (American Choral Director's Association), MENC (Music Educator's National Conference) and AGO (American Guild of Organists).
Tickets are $20 for General Admission, $15 for Friends of the Sanctuary, and only $5 for school and college students with ID. Tickets may be purchased online below by selecting your ticket option and clicking on the "Buy Now" link to use credit or debit card or your PayPal account. If ordering two or three ticket types, each type must be purchased in a separate order. You will receive an email confirmation of each purchase. Online tickets will be held at the box office; please print and bring your receipt in case it is needed to confirm your purchase.
GROUP RATES are now available! Choose the Group ticket option and each ticket will have a $5 discount from the General Admission price (same as the Friends of the 1867 Sanctuary ticket price)! Group rate tickets must be purchased in quantities of 10 or more within a single purchase. Group rates apply only to advance online sales and cannot be purchased at the box office prior to the concert. Groups must present themselves at the box office at the same time. For group orders of fewer than 10 people, the full General Admission rate (less the amount prepaid) may be required at the box office at the time of admission. Group rates do not apply to Student prices.
Tickets may be purchased at the door as space allows using cash, check or credit card.
The 1867 Sanctuary is wheelchair accessible using the ramp on the left side of the building.
For only $30.00 per year ($50.00 for families), members of the Friends of the 1867 Sanctuary receive advance notice of special events, our member newsletter, gift shop discounts and $5 off our General Admission concert tickets! BECOME A MEMBER NOW!
Return to Grand Opening Festival Calendar
October 19, 2016
Copyright © 2016 · All Rights Reserved · Preservation New Jersey


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I now have all the audio I need for Homemade Nonsense Volume III. The podcast is going to tell the story of what exactly happened when I did my final comedy gig. This one is the last one because the nature of my workload is about to change quite dramatically so I won't realistically be able to keep up the stand up. It'll have to take a backseat for a bit. Shame, but there we are.
So, as a consequence I'm going to leave off news of exactly what happened at my final ever Beat The Frog and put it all in the podcast.
How exciting.
See how I'm forcing my readers to download it? They have no choice.
Don't forget you can still get the half an hour long chunk of the phone in show I made available a bit back. Go there now and download it. It's great. You'll love it.
I look forward to explaining why exactly I won't be doing stand up anymore very soon on this blog.


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To characterize scleral biomechanics in both eyes of eight monkeys in which chronic intraocular pressure (IOP) elevation was induced in one eye.
Each posterior sclera was mounted on a pressurization apparatus, IOP was elevated from 5 to 45 mm Hg while the 3D displacements of the scleral surface were measured by speckle interferometry. Finite element (FE) models of each scleral shell were constructed that incorporated stretch-induced stiffening and multidirectionality of the collagen fibers. FE model predictions were then iteratively matched to experimental displacements to extract unique sets of scleral biomechanical properties.
For all eyes, the posterior sclera exhibited inhomogeneous, anisotropic, nonlinear biomechanical behavior. Biomechanical changes caused by chronic IOP elevation were complex and specific to each subject. Specifically: (1) Glaucomatous eyes in which the contralateral normal eyes displayed large modulus or thickness were less prone to biomechanical changes; (2) glaucomatous scleral modulus associated with an IOP of 10 mm Hg decreased (when compared with that of the contralateral normal) after minimal chronic IOP elevation; (3) glaucomatous scleral modulus associated with IOPs of 30 and 45 mm Hg increased (when compared with that of the contralateral normal) after moderate IOP elevation; and (4) FE-based estimates of collagen fiber orientation demonstrated no change in the glaucomatous eyes.
Significant stiffening of the sclera follows exposure to moderate IOP elevations in most eyes. Scleral hypercompliance may precede stiffening or be a unique response to minimal chronic IOP elevation in some eyes. These biomechanical changes are likely to be the result of scleral extracellular matrix remodeling.
Glaucoma is the second leading cause of blindness worldwide1,2 and leads to vision loss through irreversible damage to retinal ganglion cell axons as they pass through the scleral canal at the optic nerve head (ONH). Although glaucoma pathogenesis is not well understood and could also involve direct damage to the retinal ganglion cells or lateral geniculate, the biomechanical environment of the ONH has been hypothesized to play an important role in the neuropathy. The sclera is an important factor in ONH biomechanics, and recent work strongly suggests that the biomechanics of the posterior sclera and lamina cribrosa are tightly coupled.3–6 The sclera is the stiffest tissue of the eye and provides the mechanical boundary conditions for the lamina cribrosa at its insertion into the scleral canal wall. Computational models have shown that scleral stiffness7 and scleral collagen fiber organization3,4 dictate the IOP-induced deformation exhibited by the ONH. Of note, the sclera exhibits significant biomechanical changes with age8 and after exposure to chronic IOP elevation, as shown herein, both of which are major risk factors in glaucoma.
To investigate scleral biomechanics as a factor in glaucomatous damage, it is important to quantify its mechanical response to IOP. The stresses and strains in the sclera are significant at normal IOPs,8,9 and exposure to elevated IOP could lead to changes in scleral biomechanics that precede the onset of glaucoma or occur very early in disease progression. We therefore focused this work on the scleral changes that are present at the early stages of the disease in nonhuman primates, which could indicate that scleral biomechanics and its IOP-induced changes contribute to both the individual susceptibility to glaucoma and its pathogenesis.
We recently described our ex vivo method to experimentally measure the 3-D deformation pattern and thickness of posterior sclera from both eyes of four young and four old monkeys after IOP elevation from 5 to 45 mm Hg.8 In that report, we modeled the posterior sclera as a nonlinear, anisotropic, inhomogeneous soft-tissue using a fiber-reinforced constitutive theory that included stretch-induced stiffening and multidirectionality of the collagen fibers.3 We then derived eye-specific sets of scleral biomechanical properties based on the experimental observations using an inverse finite element (FE) method.9 Our results showed that posterior and peripapillary sclera in old monkeys is significantly stiffer than that from young monkeys and is therefore subject to higher stress but lower strain at all levels of IOP. Whether age-related changes in scleral biomechanics contribute to the development and progression of glaucoma remains to be determined.
We have demonstrated significant alterations in the architecture of the laminar and peripapillary scleral connective tissues at the earliest stage of experimental glaucoma (EG) in the monkey eye.10–18 In the lamina cribrosa of EG monkey eyes, permanent deformation and thickening,16 accompanied by increases in both total connective tissue volume and total number of laminar beams, were observed.15 In addition, we observed posterior deformation accompanied by thinning in the peripapillary sclera in the EG eyes.13,14 These morphologic changes are likely to result from both physical deformation in response to the applied load (IOP) and connective tissue remodeling in response to chronically elevated stress and strain levels.15,19,20 Glaucomatous remodeling of the connective tissues by optic nerve head (ONH) astrocytes and scleral fibroblasts may in turn contribute to retinal ganglion cell axonal damage through direct and/or indirect cellular mechanisms.21,22
We have shown that the biomechanical properties of the peripapillary sclera are altered in early EG using uniaxial testing and linear viscoelastic theory.11 That study demonstrated a significant increase in the equilibrium modulus of the peripapillary sclera from glaucomatous monkey eyes, but no changes were seen in the time-dependent viscoelastic parameters in those eyes.
In the present study, we used novel three-dimensional (3-D) experimental and computational techniques3,8,9 to characterize peripapillary and posterior scleral biomechanics in both eyes of eight adult monkeys in which one eye had been exposed to chronic, laser-induced IOP elevations of modest to substantial magnitude and duration.
All experiments adhered to the ARVO Statement for the Use of Animals in Ophthalmic and Vision Research. In this study, EG was induced in one eye of eight rhesus monkeys that had no initial ocular abnormality. Our protocol for generating and detecting laser-induced EG has been fully described in previous reports.12–14,16 Briefly, both eyes of each monkey underwent IOP measurement (Tonopen XL; Reichert, Depew, NY) and confocal scanning laser tomographic (CSLT) imaging (Heidelberg Retinal Tomograph; Heidelberg Engineering, Heidelberg, Germany) at manometrically controlled IOP of 10 mm Hg on three to five occasions while normal. Then, imaging was performed every 1 to 2 weeks after laser photocoagulation of the trabecular meshwork (OcuLight TX; Iridex, Mountain View, CA) in one eye, until the onset and/or progression of CSLT-detected ONH surface change by mean position of the disc (MPD) or HRT II topographic change analysis (TCA).23 Four monkeys were observed until the onset of MPD change (confirmed twice), which is our existing criterion for onset of early experimental glaucomatous damage to the ONH.24 Four additional monkeys were observed up to MPD onset and at least one episode of further progression (confirmed twice) to achieve a more advanced stage of ONH (and presumably scleral) alteration. All the monkeys were killed after an intramuscular injection of ketamine/xylazine with an intravenous injection of pentobarbital sodium.
The load-bearing connective tissues of the posterior sclera directly experience glaucomatous alterations of intraocular pressure,11 whereas they are likely to be only minimally affected by glaucomatous alterations to the neural and connective tissues of the ONH. We therefore chose to stage the animals by EG eye cumulative IOP insult rather than by the magnitude of MPD or TCA ONH surface change. We defined cumulative IOP insult for each EG eye to be the difference between the area (computed using the trapezoidal rule) under its IOP-time curve (Fig. 1, red) minus that of its contralateral normal eye (Fig. 1, blue;). Monkey 1 therefore received the lowest cumulative IOP insult and monkey 8, the highest (Table 1).
For this study, we used an experimental protocol for pressure testing the posterior scleral shell that has been fully described in our other reports.8,9 Briefly, all monkeys were anesthetized with an intramuscular injection of ketamine/xylazine and killed with an intravenous injection of sodium pentobarbital. The eyes were enucleated immediately after death, and the posterior scleral shells were cleaned of intra- and extraorbital tissues and individually mounted on a custom-built pressurization apparatus (see Ref. 9 for a detailed description of the apparatus). After mounting, each shell was blotted dry, covered with a white contrast medium (ProCAD Contrast Medium; Ivoclar, Schaan, Lichtenstein), and immediately immersed in isotonic saline at room temperature. IOP was incrementally increased from 5 to 45 mm Hg, and the resulting full-field 3-D displacements of each shell surface were recorded at a resolution of 0.1 μm with an electronic speckle pattern interferometry sensor25,26 (Q100; Ettemeyer AG, Ulm, Germany). Internal pressure was then reset to 5 mm Hg, and scleral topography was manually measured with a 3-D digitizing arm (MicroScribe G2X; Immersion, San Jose, CA). Scleral thickness was also measured at 20 predetermined locations (see Ref. 9 for details) with a 20-MHz ultrasound transducer (PacScan 300P; Sonomed, Inc., Lake Success, NY). The topography and thickness data were combined to create an eye-specific reference geometry for the posterior scleral shell in each subject.
Our constitutive theory for simulating the biomechanical behavior of the posterior sclera and our inverse FE method of extracting mechanical properties based on the scleral pressure testing outlined above have been fully described in our previous reports.3,8,9 Both are briefly explained as follows.
We modeled the posterior sclera as a nonlinear, anisotropic, inhomogeneous soft tissue by using fiber-reinforced composite theory. Within this framework, we assume that scleral collagen fibers provide most of the resistance to IOP and are responsible for two important biomechanical characteristics (nonlinearity, described with the parameters c3 and c4, and anisotropy, with θp and k; Fig. 2). The model assumes that scleral collagen fibers are embedded within a ground substance matrix, not to be confused with the extracellular matrix, which includes all the remaining tissue constituents (e.g., water, fibroblasts, proteoglycans, and elastin). Hence, the ground substance matrix is of low stiffness compared with the collagen fibers, and its mechanical behavior is characterized with a single intrinsic parameter, c1, as isotropy is assumed (i.e., no preferred orientation).
Using our inverse FE method, we estimated a set of thirteen model parameters (c1, c3, c4, k1, k2, and θp1 to θp8; Fig. 3) from the scleral mechanical testing data for each posterior scleral shell. This strategy uses a genetic optimization algorithm (differential evolution47–49) to generate a unique value for each of the 13 model parameters by simultaneously varying all 13 parameters until the FE-predicted displacements closely match the experimental deformations at five IOP levels simultaneously (7, 10, 20, 30, and 45 mm Hg).
Two additional measures of scleral stiffness were derived from the 13 output parameters: the tangent modulus and the structural stiffness. Because of their high clinical relevance, these two stiffness measures are emphasized in the Results and Discussion sections.
The model parameters c3 and c4 describe the stretch-induced stiffening of the collagen fibers, but do not represent a direct measure of scleral stiffness as a function of IOP. To capture scleral stiffness, the tangent modulus along the preferred fiber orientation was calculated and mapped. The tangent modulus is an estimate of local scleral material stiffness, varies with IOP, and is independent of geometry.9
Clinical discussions concerning the observable mechanical behavior of the corneoscleral shell often confuse the difference between the biomechanical material properties of the tissue (the stiffness of the bulk material itself), its architecture (thickness), and the resultant net structural behavior. Although the material properties of the tissues are intrinsic to the material itself, the structural stiffness of the shell incorporates both the material properties and geometry of a complex load-bearing structure into a composite measure of the structure's resistance to deformation. We defined the structural stiffness of the sclera as the local product of the scleral thickness and the tangent modulus to capture the mechanical behavior of the scleral shell as a structure in response to acute IOP elevations.
Regional thickness differences between the normal and glaucomatous eyes of each animal were compared to the regional thickness differences between bilaterally normal eyes of eight previously studied monkeys,8 by using a multiple linear regression.
Posterior displacements between the two eyes of each monkey could not be directly compared, because all displacements were computed with respect to the position of the scleral clamp, which varied to a certain degree among eyes. However, the derived output measures (stress, strain, and structural stiffness) can be directly compared because they are unaffected by scleral clamp position.
Tangent modulus and structural stiffness distributions for the eight adult normal eyes were compared to the previously reported data for four young and four old bilaterally normal monkeys at IOPs of 10, 30, and 45 mm Hg using the general estimating equation method as used in our previous report.8
Regional distributions of maximum principal stress, maximum principal strain, tangent modulus, and structural stiffness were compared between the normal and glaucomatous eyes of each monkey at IOPs of 10, 30, and 45 mm Hg by paired t-tests. Note that the regional data for these output measures (i.e., stress, strain, modulus, and stiffness) were not normally distributed, which violates the assumptions underlying t-tests. To address this, and as commonly performed using the central limit theorem, 3000 means were calculated for each parameter, each consisting of 300 randomly sampled values from the original distribution. The values of 300 and 3000 were chosen as the minimum values necessary for the t-statistic values to be unaffected by non-normality. These normally distributed means were then used in the t-tests to compare the parameters for normal and EG eyes. For each output measure (i.e., stress, strain, modulus, and stiffness), alterations due to chronic IOP elevation were deemed to be significant for that particular monkey when the t statistic from comparing the normal and glaucomatous eye of that monkey exceeded all t statistics computed from comparisons of bilaterally normal eyes of eight other monkeys.8
Finally, anisotropy parameters (i.e., preferred fiber orientation and fiber concentration factor) were compared between groups by paired t-tests in both the peripapillary and peripheral sclera.
Scleral thickness, experimentally measured at an IOP of 5 mm Hg at 20 locations and interpolated over each scleral shell, is shown in the first row of Figure 4 and in Figure 5 (averaged values). In all eyes, the peripapillary sclera was much thicker (276–498 μm on average in the normal eyes) than the peripheral sclera (157–252 μm on average in the normal eyes; P < 0.0001). Among those regions considered (inferior, inferonasal, nasal, superonasal, superior, superotemporal, temporal, and inferotemporal), the sclera was significantly thicker in the temporal region for all 16 eyes (P < 0.001).
For each eye, good agreement was obtained between the FE-predicted and experimentally measured posterior displacements between 5 and 30 mm Hg (second and third rows of Fig. 4; Table 1; maximum averaged nodal error, 4.08 μm). Although not all cases are included in Figure 4, good agreement was also obtained for each IOP range (5–7, 5–10, 5–20, 5–30, and 5–45 mm Hg) and for all displacement components, suggesting that our constitutive model was able to accurately capture the complex 3-D deformations of the monkey peripapillary and posterior sclera after acute elevation of IOP.
For each eye, the sclera stiffened considerably as IOP increased, as demonstrated by the large increase in tangent modulus in Figure 6 (2.2–7.7-fold increase from 10 to 45 mm Hg in the peripapillary sclera from normal eyes), and also by the large increase in structural stiffness as IOP increased (Fig. 7). This phenomenon is also demonstrated by the nonlinear relationship between maximum principal strain and IOP (Fig. 8). Note that maximum principal strain values were consistently low in the normal eyes, as shown in blue in Figure 8, and varied between 0.32% and 1.49% on average in the peripapillary sclera at 45 mm Hg. Conversely, maximum principal stress increased linearly with IOP (Fig. 8) and was considerably higher than IOP (16 to 30 × IOP on average in the peripapillary sclera from normal eyes at all IOP levels).
For each eye, the sclera was highly inhomogeneous as demonstrated by the regional variations in tangent modulus (Fig. 4, row 4). On average, the tangent modulus was higher in the peripheral sclera (17–60 MPa in the normal eyes) than in the peripapillary sclera (15–50 MPa in the normal eyes).
The sclera was highly anisotropic in all eyes as demonstrated by the non-zero fiber concentration factors shown in Table 1 and Figure 2. The fiber concentration factor in the peripapillary sclera (k2) was higher than that in the peripheral sclera (k1) in 10 of the 16 eyes studied (Table 1). Maps of the predicted preferred fiber orientation for all eight regions of each eye are shown in Figure 4, where (white) indicates a preferred fiber orientation perpendicular to the scleral canal (meridional, θp = 0°), and // (black) indicates a preferred fiber orientation tangent to the scleral canal (circumferential, θp = 90°). Figure 9 shows the pooled distributions of the preferred fiber orientation for all 16 eyes in both the peripapillary and peripheral sclera. In general, the mean preferred fiber orientation for all tested eyes was tangent to the scleral canal in both scleral regions, but this effect was more pronounced in the peripapillary sclera.
We considered all eight normal eyes from this study as being part of one age group (adult, 17.3 ± 5.1 years). Note that the monkey lifespan is about one third the human lifespan.51 We compared tangent modulus and structural stiffness distributions from the monkeys in this study to that of two other age groups (young, 1.5 ± 0.7 years; old, 22.9 ± 5.3 years) from our previous report.8 Using the generalized estimating equation method, we found that the tangent modulus and structural stiffness in the peripapillary sclera significantly increased with age (Table 2). Using a multiple linear regression, we found that eyes from old monkeys were significantly thinner than eyes from adult monkeys (P < 0.05) but found no significant differences in thickness between eyes of young and adult monkeys (P > 0.05). Peripapillary scleral thickness averaged 389 ± 53 μm in young eyes,8 381 ± 47 μm in adult eyes (this study), and 349 ± 49 μm in old eyes.8
We observed no significant changes (exceeding the physiologic intereye difference of normal monkeys) in tangent modulus (Fig. 6) and structural stiffness (Fig. 7) between the normal and glaucomatous eyes of monkeys 2, 3, and 8.
We observed significant changes in tangent modulus (Fig. 6) and structural stiffness (Fig. 7) between the normal and glaucomatous eyes of monkeys 1, 4, 5, 6, and 7. The glaucomatous eye in monkey 1 exhibited a significant decrease in tangent modulus (10 mm Hg) and structural stiffness (10 and 30 mm Hg). Note that monkey 1 received the lowest cumulative IOP insult and these changes manifest at lower IOPs. The glaucomatous eyes in monkeys 4, 5, and 6 exhibited a significant increase in tangent modulus and structural stiffness at higher IOPs (30 and 45 mm Hg). Monkey 7 exhibited a significant increase in tangent modulus at 45 mm Hg in the peripheral sclera only.
We observed significant changes (exceeding the physiologic intereye difference of normal monkeys) in maximum principal strain in two of the eight monkeys (Fig. 8, 1 and 4) and in maximum principal stress in two of the eight monkeys (Fig. 8; 4 and 6).
We did not detect any changes in the fiber concentration factor (P > 0.05) or in the preferred fiber orientation (P > 0.05) in either the peripapillary or peripheral regions between the normal and glaucomatous eyes.
Finally, peripapillary and peripheral scleral thickness in both the normal and glaucomatous eyes of each monkey are shown in Figure 5. Although a trend toward scleral thinning was observed in the glaucomatous monkey eyes, overall and regional change in scleral thickness between glaucomatous and normal eyes was insignificant when compared to the physiologic intereye difference from eight normal monkey eyes (P > 0.05).
In this study, we characterized scleral biomechanical behavior in both eyes of eight adult monkeys in which one eye had been exposed to chronic, laser-induced IOP elevations of modest to substantial magnitude and duration. The following are the principal findings of this report.
When considered as a group of 16 monkey eyes without regard for experimental condition (normal versus EG) the eyes in this report behaved similarly to the eyes from bilaterally normal monkeys from a previous report,8 as follows. First, the monkey posterior sclera is thicker in the peripapillary and temporal regions. Second, the monkey posterior sclera is a nonlinear, anisotropic, and inhomogeneous soft tissue. Third, concentrations of tangent modulus, structural stiffness, and maximum principal stress and strain were evident adjacent to the scleral canal in each eye. Fourth, maximum principal stress magnitude was substantially higher than IOP. Fifth, maximum principal strain magnitude was low. A detailed discussion of these results can be found in our previous report.8
Scleral biomechanical properties (i.e., tangent modulus and structural stiffness; Table 2) in the eight normal adult eyes (mean age, 17.3 ± 5.1 years) from this study fell between those from young (1.5 ± 0.7 years) and old (22.9 ± 5.3 years) monkeys from our previous aging study.8 These findings separately confirm our initial report of age-related stiffening of the posterior sclera by establishing that adult eyes are stiffer than young ones but less stiff than old eyes. This result is likely to be related to increasing scleral cross-links with age28,52; details on this topic can be found in our previous report.8
We have reported a decrease in scleral thickness in old compared with young normal monkey eyes. In that study, we found that adult monkey sclera was not significantly thinner than young monkey sclera, but was significantly thicker than sclera from old monkey eyes. This finding is important because it suggests that the rate of scleral thinning, if present, may accelerate with age. Whether scleral thinning with age could accelerate the progression of glaucoma in elderly patients remains unclear. In monkeys, we observed a reduction in peripapillary scleral thickness of approximately 8.4% from adult to older eyes on average. In comparison, we observed an increase in tangent modulus of 37.4% (at 30 mm Hg). If both tangent modulus and thickness contribute similarly to the overall stiffness of the scleral shell (i.e., structural stiffness), then age-related changes in peripapillary scleral thickness will have a lesser impact on IOP-induced deformation than those of the tangent modulus. The relationship between glaucomatous susceptibility and scleral stiffness is likely to be complex, individual specific, and involve many interacting factors.
The major findings were as follows: First, significant changes in tangent modulus and structural stiffness were observed in some monkeys between the normal eyes and those exposed to chronic IOP elevation. Second, changes in FE-based scleral anisotropy measures were not observed between normal and glaucomatous monkey eyes. Finally, although all glaucomatous eyes exhibited thinner peripapillary sclera than their contralateral normal eyes (Fig. 5), these differences did not achieve statistical significance.
We observed significant changes in scleral tangent modulus (Fig. 6) and structural stiffness (Fig. 7) between the normal and glaucomatous eyes of monkeys 1, 4, 5, 6, and 7. The glaucomatous eye in monkey 1 exhibited a smaller tangent modulus at 10 mm Hg (Fig. 6) and structural stiffness at 10 and 30 mm Hg (Fig. 7) than those of its contralateral normal. Burgoyne et al.53 have reported hypercompliance of the ONH surface within 4 to 8 weeks of chronic IOP elevation in monkeys and suggested that weakening of the load-bearing structures of the eye (lamina cribrosa and peripapillary sclera) through connective tissue damage might be responsible for this phenomenon at the earliest stage of glaucomatous optic neuropathy. Moreover, a computational modeling study by Downs et al.54 reported weakening of the lamina cribrosa structure at the earliest stage of glaucomatous optic neuropathy. It is interesting to note that weakening of the sclera was only present in the glaucomatous eye of monkey 1, the eye that received the smallest cumulative IOP insult (432 mm Hg × days) and had the lowest maximum measured IOP (24 mm Hg). It should also be noted that the decrease in tangent modulus in the glaucomatous eye of monkey 1 was only significant at low IOP (10 mm Hg).
In 1957, Roach and Burton55 studied the roles of elastin and collagen in arteries. They found that elastin alone was responsible for resisting pressure-induced arterial deformations at low pressure and that collagen alone was responsible for resisting pressure-induced arterial deformations at high pressure. These results were recently confirmed by Fonck et al.56 Based on these studies, it could be that only damage to elastin occurred in the glaucomatous eye of monkey 1, and this represents the earliest stage of scleral damage and/or remodeling in glaucomatous progression.
We observed a significant increase in scleral tangent modulus at IOPs of 30 and 45 mm Hg in both the peripapillary and peripheral regions in monkeys 4, 5, and 6, and a significant increase in scleral tangent modulus at 45 mm Hg in the peripapillary region in monkey 7 (Fig. 6). The glaucomatous eyes in monkeys 4, 5, and 6 also exhibited larger structural stiffness in the peripapillary sclera at IOPs of 30 and 45 mm Hg (Fig. 7). These findings suggest that at higher levels of cumulative IOP insult, the posterior sclera stiffens through extracellular matrix remodeling. This finding is consistent with that in our previous uniaxial study of scleral biomechanics in glaucomatous monkey eyes11 and may be a protective mechanism that limits scleral canal expansion and related deformation of the ONH at elevated IOPs. Zeimer and Ogura57 reported that human ONHs became more rigid with severe glaucoma, a result that is consistent with this hypothesis. Roach and Burton55 also demonstrated that collagen alone was responsible for limiting arterial deformations at high pressure, which suggests that the observed increase in tangent modulus in glaucomatous eyes at high IOPs may be due primarily to collagen remodeling.
Even though tangent modulus changes occurred in five monkeys (1, 4, 5, 6, and 7), significant changes in maximum principal strains (monkeys 1 and 4) and maximum principal stress (monkeys 4 and 6, in the peripapillary sclera only) were each observed in only two monkeys. This result suggests that overall changes in scleral thickness (generally lower in the glaucoma eyes) and tangent modulus (generally higher in the glaucoma eyes) at this stage of damage interact to maintain the stress and strain ranges typical of sclera in normal eyes.
In this study, we observed no significant changes in either scleral tangent modulus (Fig. 6) or structural stiffness (Fig. 7) between the normal and glaucomatous eyes in three monkeys (2, 3, and 8). Since monkey 8 received the highest cumulative IOP insult (7714 mm Hg × days), we would have expected to observe the largest changes in the glaucomatous eye of this monkey. Of note, at IOPs of 30 and 45 mm Hg, the normal eyes of monkeys 3 and 8 had the two highest mean tangent moduli and structural stiffnesses in the peripapillary sclera. Assuming that before IOP insult, the glaucomatous eyes in monkeys 3 and 8 were as stiff as their contralateral normals, these results suggest that stiffer scleral shells may be less prone to biomechanical changes when exposed to chronic IOP elevation, presumably because they are more resistant to IOP-induced strain.
Several studies have reported that mechanical strain applied to scleral fibroblasts can trigger the release of MMPs and TIMPs,58–61 which can then lead to remodeling of the scleral extracellular matrix. IOP-induced scleral shell expansion increases scleral strain in all eyes, but in eyes with the stiffest sclera, these increases may be too small to elicit a fibroblast response. Note that mean strain levels at 45 mm Hg in monkeys 3 and 8 were 0.30% and 0.32% respectively, which were smaller than all the reported values in the aforementioned in vitro studies on scleral fibroblast response to strain (a minimum of 0.45%58).
In the case of the third monkey, which demonstrated no change in tangent modulus or structural stiffness (monkey 2), the normal eye had the most scleral thickness (Fig. 5), and lowest tangent modulus (Fig. 6) among all normal eyes. Its mean structural stiffness was therefore comparable with the mean structural stiffness of all normal eyes. These data suggest that mean structural stiffness may not always predict which eyes will undergo IOP-induced scleral remodeling and that scleral thickness may also be an important contributor in predicting which eyes will be less sensitive to biomechanical change.5,46
In this study, we did not observe significant differences in predicted scleral anisotropy (preferred fiber orientation and fiber concentration factor) between normal and glaucomatous monkey eyes. The preservation of anisotropy was recently reported for the remodeled monkey lamina cribrosa in EG,15 suggesting that remodeling of the peripapillary and posterior sclera must be accompanied by changes in elastin/collagen content and/or fiber diameter, but not fiber reorientation. This is consistent with the work of Quigley et al.,62 who reported glaucomatous changes in collagen and elastin content and collagen fiber diameter in the peripapillary sclera in human and monkey eyes.
Although we did observe a trend toward scleral thinning in the glaucomatous eyes, this difference did not exceed the physiologic intereye differences in the eight normal monkeys. Using traditional histology, we previously reported significant thinning of the posterior10 and peripapillary sclera13,14 in monkey eyes subjected to chronic IOP elevation. Scleral morphologic changes should be assessed in future studies to understand whether scleral thinning due to chronic IOP elevation is a consequence of strain-induced fluid exudation, tissue loss, or a combination of the two.
Overall, our data suggest that a stiffer scleral shell is biomechanically beneficial and is less apt to remodel in response to elevated IOP. In normal eyes, we suggest that the sclera shields the ONH from biomechanical insult by resisting large overall deformation (through uncrimping and stiffening of the collagen fibers: nonlinearity) and large scleral canal expansion (through the presence of a circumferential peripapillary ring of collagen fibers: anisotropy).3,4,8 We hypothesize that the scleral stiffening in response to moderate exposure to chronic IOP elevations shown herein is an effort to maintain the biomechanical homeostasis of the scleral shell and indirectly, the ONH. As such, this response could act to slow glaucomatous progression in eyes with higher IOPs and/or more compliant scleral shells. Surprisingly, the argument that a stiff scleral shell shields the ONH from glaucomatous damage seems to contradict the facts that (1) the sclera stiffens with age.8 and (2) the elderly are more susceptible to glaucoma.63 To reconcile all these observations, it should be noted that collagen fibers become brittle with age28 and therefore it is plausible to suggest that scleral stiffening with age is an unhealthy mechanism as opposed to scleral stiffening because of the remodeling seen in this study. In addition, there are other factors that are likely to contribute to glaucomatous damage, including blood flow and cellular reactivity, and these factors change with age as well. We also caution the reader that stiffness is a broad term, and reporting a single stiffness value for the sclera does not represent its biomechanical response well. Scleral stiffness is a function of IOP (nonlinearity), orientation (anisotropy), and location (heterogeneity). This complexity should be taken into account when evaluating the contribution of scleral biomechanics to glaucoma pathogenesis.
Several limitations should be recognized when viewing this work. Although the general limitations of the method have been discussed at length in our previous reports,3,8,9 we will briefly revisit them and then discuss those inherent to this study.
First, IOP was measured in both eyes of each monkey over time (Fig. 1), but we did not characterize the diurnal and nocturnal IOP fluctuations that are known to exist in monkeys. Diurnal IOP fluctuations have been shown to be large in monkeys, especially in glaucomatous eyes, in which they are highly individual-specific and can exceed 10 mm Hg.64 Therefore, our definition of cumulative IOP insult (change in area under the IOP/time curve) should be regarded with caution and was used here only to provide a rough estimate of the cumulative insult from IOP. An implantable telemetric IOP transducer is currently being developed for monkeys that should allow for continuous measurements of diurnal and nocturnal IOP fluctuations and more accurate characterization of IOP insult (Downs JC, et al. IOVS 2008;49:ARVO E-Abstract 2043).
Second, because the sclera is approximately 90% collagen by weight,65 we assumed that collagen is the primary fibrous element66 of this tissue. We lumped all other tissue constituents into the ground substance matrix, a common practice in soft-tissue FE modeling.4,30,67–71 Because our study suggests that both elastin and collagen play important roles in glaucomatous damage, further experimental and theoretical work is needed to address how these components separately contribute to scleral biomechanics.
Third, we did not characterize scleral biomechanics between 0 and 5 mm Hg in the tested eyes, because the scleral shells required an IOP of approximately 4 mm Hg to sustain their shape. IOPs in the range of 0 to 5 mm Hg are rarely measured in monkey eyes, and so ignoring the initial IOP loading should not compromise the clinical importance of our results. We are developing methods that will allow the initial scleral stress and strain that exist in the tissues at 5 mm Hg to be estimated and included in future studies.
Fourth, the ONH tissues were assumed to be linearly elastic, with a common elastic modulus of 1 MPa assigned to the ONH region for both normal and glaucomatous monkey eyes. Our previous sensitivity study showed that ONH elastic moduli in the range of 0.1 to 5 MPa did not affect the results of our scleral biomechanical property fitting9; hence, our assumption should not affect the results presented herein.
Fifth, within our models the ONH was represented as an elliptical cylinder, with its external boundary located at the outer aspect of the dural sheath insertion (as measured with the 3-D digitizer) and therefore included regions of the immediate peripapillary sclera and scleral flange. As a result of this anatomic convention, we could not estimate biomechanical properties for that region of the sclera that lies immediately under the dural sheath insertion.
Sixth, we divided each reconstructed scleral shell into eight regions to assess regional variations in predicted collagen fiber orientation (θp1–θp8). This division was made because a preliminary study confirmed that a combination of eight regions and 13 model parameters was the minimum requirement to obtain good agreement between the FE-predicted and experimentally measured displacements (Fig. 4), while also ensuring a unique set of model parameters. Future studies will include a more rigorous assessment of regional variability.
Seventh, all experiments were conducted at room temperature (22°C) because the incorporation of a heated saline chamber at 37°C proved incompatible with the speckle interferometry sensor we used. Because all experiments were consistently performed at the same temperature, all hypotheses tested regarding the changes in scleral biomechanics should be valid.
Eighth, collagen fiber alignment was not measured directly, but was predicted using the models' fits to experimental scleral deformations (Fig. 4). Because of the relatively coarse discretization of the regions (discussed above) we could only report an overall approximation of the preferred collagen fiber orientation within each region. Future inclusion of experimental measures of collagen fiber orientation as input parameters, as currently being measured with small-angle light-scattering (Girard MJ, et al. IOVS 2010;51:ARVO E-Abstract 2128), will allow for finer regional discretization for the remaining fitted parameters and improve our characterization of scleral anisotropy.
Finally, seven of the eight monkeys (monkeys 2–8) began this study while the investigators laboratories were still located at the LSU Eye Center in New Orleans, and were unavailable for study for more than 3 months after Hurricane Katrina. Monkeys 2 and 3 and 6 to 8, however, had already undergone initial laser, and so the accuracy of their cumulative IOP exposure calculations is affected by the lack of IOP data between 30 August and 15 December 2005, (Fig. 1, shaded area) and the diminished frequency of testing after their arrival at Devers. In addition, these monkeys (2, 3, and 6 to 8) were examined only sporadically until the laboratory at Devers became fully operational. At the time of the hurricane, monkeys 4 and 5 had not yet been lasered, and so the frequency of IOP measurement and assessment of pre- and postlaser cumulative IOP exposure in these animals was unaffected by the hurricane. Monkey 1 was purchased at Devers and was altogether unaffected by these events.
In summary, new experimental and computational methodologies3,9 were used in this study to characterize scleral biomechanics in normal and glaucomatous monkey eyes. These methods have broad applicability and can be applied to other thin, soft tissues with multidirectional collagen fibers. The long-term goal of this work is to establish the pathophysiologic mechanisms linking connective tissue stress and strain within the peripapillary sclera and lamina cribrosa to retinal ganglion cell axon damage and death. Given the important role of scleral biomechanics in determining the biomechanics of the ONH, new strategies for the clinical visualization and measurement of peripapillary and posterior scleral biomechanics are needed.
Overall, our results suggest that significant changes in the biomechanical behavior of the sclera are associated with exposure to chronic IOP elevations. These changes are complex and individual-specific and are likely to be the result of scleral extracellular matrix remodeling. Our findings suggest that (1) scleras with initially large tangent modulus or thickness are more resistant to IOP-induced biomechanical changes, (2) a decrease in tangent modulus (possibly due to elastin damage) occurs at the earliest stage of glaucoma, (3) an increase in tangent modulus (possibly associated with collagen changes) occurs at moderate stages of glaucoma, and (4) collagen fiber reorientation does not occur on a large scale in glaucomatous eyes. The sclera drives the levels of stress and strain transmitted to the ONH, so changes in scleral biomechanical behavior will inevitably perturb the biomechanics of the ONH.7 Scleral remodeling induced by exposure to chronic IOP elevations may provide a protective mechanism for the contained ONH and act to keep stress and strain levels within the normal range, but future work is necessary to establish the link between scleral/ONH biomechanics and glaucomatous vision loss.72–74
The authors thank Juan Reynaud, Jonathan Grimm, Wenxia Wang, and Stuart Gardiner for their contributions to this work.
Presented in part at the annual meeting of the Association for Research in Vision and Ophthalmology, Fort Lauderdale, Florida, May 2009.
Supported by NIH Grant R01EY11610 (CFB) and the Legacy Good Samaritan Foundation, Portland, OR.
Disclosure: M.J.A. Girard, None; J.-K.F. Suh, None; M. Bottlang, None; C.F. Burgoyne, None; J.C. Downs, None
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Pain in throat
I started to get pain in throat which had increased from evening. I took betadine n gargled twice but no relief.
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Hello there. I can understand your concern. It could be pharyngitis. Do betadine throat gargles thrice a day and drink warm water and it usually subsides within a day or two. If it still persists, then you would need antibiotics so consult a general physician in person for further evaluation and management. Best regards. Dr. Anil kumar jain. Bangalore.
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Apple is being pressured to release $76 Billion Cash
Investors are begging Apple to pay dividends from its huge stash of cash. With billions in reserve for that rainy day, Apple is playing the game the same way Steve Jobs would have had them play it. This is the fortitude that took them to the top.
Finance chief Peter Oppenheimer is refusing to release a dime in dividends. This same request has been made ad nauseum, but Oppenheimer, in the spriti of Jobs, is keeping the cash for the next opportunities to arise. With stocks twice as high as the previous year, Oppenheimer made the pointed argument that they have this cash exactly because of this mentality. The cash and investments Apple holds are worth $76.2 billion.
The call to open the coffers comes only days after the death of Steve Jobs. However, he saved Apple from bankruptcy and took it to the pinnacle of the most valuable tech company in the world, and even the most valuable company in the world – beating Exxon for one week. His attitude of hoarding cash put them in the position they are now in, where they have so much money they don't know what to do with it. As a result, Oppenheimer is being assaulted with demands to make a dividend payment or a stock buyback available.
Oppenheimer holds a bachelor's degree from California Polytechnic University of San Luis Obispo, as well as an M.B.A. from the University of Santa Clara. He replaced Fred Anderson upon his retirement as CFO, in 2004. Apple had only $5 billion in cash and investments when Oppenheimer took over, but during the following three years it tripled to over $13 billion. The next three years saw another leap to over $40 billion in 2010.
Keith Goddar, CEO of Capital Advisors Inc., and the largest holder of Apple stocks, said, "It wouldn't hurt their position any if they paid out a small dividend." In 1997, Apple wasn't sure if it would be able to pay its employees. Steve Jobs took the helm at that time and now, with the advent of the iPhone and the iPad, Apple is sitting pretty. Jobs wanted to steer Apple into new market opportunities with the cash, rather than returning it to shareholders.
In spite of the stock market slump, Apple's stock has increased 15% this year alone. Apple is valued at $342.8 billion, placing it far ahead of the second largest tech company, Microsoft. In fact, Apple is 60% bigger than Microsoft today.
Other large tech companies have not had such an outlook on their cash management. For example, Microsoft and Google utilize their cash for large acquisitions. Cisco paid out dividends starting this year, following the $40-billion mark.
Apple is using some of its cash to purchase future supplies of the most significant components for their products. Capital expenditures ad components will take up about $3.9 billion in the next two years, according to Oppenheimer. This is a company that has demonstrated the dedication to good cash management principles in order to be ready for any potential strategic opportunities.
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/ Having read this I thought it was really informative. I appreciate you spending some time and energy to put this short article together. I once again find myself personally spending a significant amount of time both reading and posting comments. But so what, it was still worth it!
Ja wie? Erst die Nase lang machen und dann nix zeigen? ;.)Aber ich bin mir ebenfalls sicher, wenn du darfst, sind wir die Ersten, die es zu sehen bekommen. Freu mich für dich!


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Category Archives: Accessories
One Simple Tote-Free Reversible Tote Bag Tutorial
I recently made this camo tote for a friend's birthday. My hubby makes excellent wine, so our gift was several bottles of wine in this camo tote. Our friend loves Joe's wine and is a real outdoorsman, so this went over really well! Of course, you can use this pattern to make all sorts of reversible totes in … [CLICK TO READ MORE]
The Iris Garden Hat, a Springtime Inspiration
All the beautiful Springtime flowers blooming inspired my latest hat PDF sewing pattern, the Iris Garden Hat. This lovely wide-brimmed hat pattern comes in sizes to fit babies to adults. It's so easy to adjust with a draw-string tie in the back. The Iris Garden Hat is perfect for gardening, going to the beach or any sunny day … [CLICK TO READ MORE]
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Living Raw Food: Get the Glow with 100 More Recipes from Pure Food and Wine Hardback
by Sarma Melngailis
Picking up where the bestselling "Raw Food/Real World" left off, executive chef and partner of "Pure Food and Wine", Sarma Melngailis invites us inside the restaurant with a hundred more recipes for delectable and healthful juices, shakes, soups, simple dishes, main courses, desserts, and cocktails.
Cool down with a Cucumber-Mint Gazpacho Soup and an Heirloom Tomato Salad with Caper Emulsion.
Surprise your family with a raw Thanksgiving dinner, complete with 'Dark Meat' Portobello, 'White Meat' Large Oyster Mushrooms, Stuffing, Mashed Celeriac, Cranberries, and Brussels Sprouts.
Satisfy your sweet tooth with a Vanilla Pear Pudding and Mexican-Chocolate Brownie with Sweet Tamale, Hibiscus Cream, and Avocado Gelato.
- Format: Hardback
- Pages: 384 pages
- Publisher: HarperCollins Publishers Inc
- Publication Date: 01/06/2009
- Category: Health & wholefood cookery
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Showing 1 - 1 of 1 reviews.
Review by jfoster_sf
This book was nice in that in had a lot of information about raw food, what you should be eating, the best foods to get your protein, calcium, iron, etc, but I didn't realize when I grabbed this to read that the author was a restaurant owner and that most of the recipes are taken from the menu, and were super complicated and would take forever to make! There were some recipes that were realistic to make at home, but not enough to justify me buying this one.
Also by Sarma Melngailis


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U.S. Supreme Court To Determine Constitutionality of Gay Marriage
Photo by Ian Witlen
Today, the U.S. Supreme Court Justices have said they will determine this spring whether the Constitution requires all states to issue marriage licenses to same-sex couples and whether every U.S. state must recognize same-sex marriages performed elsewhere.
Such answers could bring marriage equality to the remaining 14 states that have bans against same-sex unions, or possibly reverse pro-gay rights' rulings that have swept the nation.
In April, the justices will review a ruling by Sixth Circuit, which upheld bans on same-sex marriage in four states: Kentucky, Michigan, Ohio and Tennessee.
In November, the federal appeals court that oversees those four states upheld their same-sex marriage bans, which reversed pro-gay rights' rulings of federal judges in each state.
In October, the justices' decision to not hear appeals supporting state bans allowed some of those lower court rulings to take effect. .
With Florida becoming the 36th state to recognize same-sex unions, the Human Rights' Campaign said that about 70 percent of Americans currently live in states where same-sex marriages are legally recognized. That's a jump from about 11 percent in 2012.
Besides the states in the Sixth Circuit, ten other states also prohibit same-sex marriages.
In Arkansas, Mississippi, Missouri, South Dakota and Texas, judges struck down anti-gay marriage laws, however the bans remain in effect pending appeals. There have been no rulings on lawsuits in Alabama, Georgia, Nebraska, and North Dakota.
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Served until 10pm
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Roasted in our smoky spice rubsticky barbeque or Louisiana hotsauce
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In slow cooked tomato sauce, just like Mamma made them
With Pietro's Italian Sausage, ground Scottish steak mince, tomato, herbs and oozing mozzarella. Served with garlic bread
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Classic sauce of egg, Parmesan, cream and garlic with smoky bacon
Classic rich beef ragu with red wine and herbs
Pasta tubes in chilli and tomato sauce Choose mild, hot or very hot. Add chicken breast £2
Pasta tubes in a rich, spicy sauce made with Pietro's Italian sausage, tomato, garlic and chilli. £2 supplement
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Starter £5.95 • Main £10.95
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Add chicken £3 • Add king prawns £4
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All burgers are made with Scottish steak mince and Pietro's Italian sausage. Served on brioche bun with skinny fries, lettuce, tomato and pickle
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Double Burger with 3 cheeses, suicide 3 chilli relish,large portion of spicy fries and garlic and chilli cheese baguette.
With vanilla ice cream
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The latest videos and audio podcasts from XXXchurch.com
January 23rd, 2014
Episode 54 of 155 episodes
Come join us in the latest episode of Open 24/7 where we spend some time talking with Jason also known as Propaganda. Then get open by visiting http://X3watch.com By xxxchurch Tags : accountability, craig gross, get open, life as a musician, open 24x7, propaganda, xxxchurch
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In a time where we're all threatened by a rhetoric of hate from the people in power; A Gay And A NonGay challenges many of our differences head on and promises that no matter who you are, or what you're into (Bruce Springsteen or Britney), love is love and gay and nongays can be friends.

An independent podcast from James Barr (@imjamesbarr) and Dan Hudson (@DanHudson). Contact us on Twitter, IG or Facebook @gaynongay


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As a junior he won four singles and eight national doubles titles before turning professional in 1987. He became a member of the British Davis Cup squad and in 1991 helped Britain regain their place in the World Group when he beat Thomas Muster in the defeat of Austria in Manchester. The same year he won the Christchurch Challenger in New Zealand, but it was not until 1994 that he broke into the world top 100, with Michael Stich and Michael Chang among his victims.
He reached three quarter-finals and two semi-finals in ATP Tour events while, at Wimbledon, he was a quarter-finalist in the mixed doubles in 1995 and reached the third round of the men's singles last year.
Petchey said: "I have had a great time in tennis and I would not have changed anything I have done in the last 10 years. Now I'm looking forward to my new career."
Pat Rafter, the No 1 seed, survived a tricky match with Martin Damm yesterday to reach today's quarter-finals of the Mercedes Cup in Los Angeles. Rafter, who defends his US Open title next month, said the knee problem that had bothered him during the French Open and Wimbledon, seemed to have been conquered as he overcame Damm 6-3, 5-7, 6-1.
He took two and a half weeks off before this event, and said that would be his last break of the season as he prepares to defend his first Grand Slam title. He was encouraged by his victory against Damm, which took one hour and 41 minutes.
Rafter's compatriot Sandon Stolle, a lucky loser who replaced the injured Thomas Enqvist in the main draw, beat another Australian, Andrew Ilie, 6-3, 7-6. Stolle will now face the fifth-seeded American, Andre Agassi, who beat Jan-Michael Gambill 6-4, 6-4.
Agassi had previously been beaten by the fast-rising American youngster with movie star looks. But this time he maintained the recent improvement in his form with a sharp display against the hard-hitting Gambill. Afterwards, he said: "You don't want to lose to someone two times in a row. It's nice to go out and play well."
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Dr. Louis Brunsting III MD Video
- Vascular Surgery Birmingham, AL
Take a minute to learn about Dr. Louis Brunsting III in this video.
Meet Doctor Louis A Brunsting. Here are his vitals: He's board certified, specializing in Thoracic Surgery with expertise in several areas. Doctor Brunsting is a highly experienced doctor, with 25 years of practice in the field Highly regarded by his patients they rate him 3 out of 4 stars. They point out his bedside manners. He has won Castle Connolly America's Top Doctors® 2006. Is affiliated with a highly rated hospital. Graduated from top rated University of Michigan Medical School. Doctor Brunsting is a published author, having articles in peer reviewed journals. His average waiting time is 32 minutes. He accepts new patients. He has a number of insurance plans accepted. Use Vitals.com to examine Doctor Louis A Brunsting from Ann Arbor, Michigan. See patient comments make an appointment or even let us help you to prepare for your visit. Vitals .com where doctors are examined.
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