But last week, Congress may have made
life less rewarding for tax exiles.
Some exiles were born and raised in
the U.S., such as John Dorrance III -- grandson of the inventor and
entrepreneur who helped found Campbell Soup Co. -- who renounced his
citizenship in 1994 and emigrated to Ireland, which has significantly lower tax
rates. Others have long lived outside the U.S. and are seeking to avoid the
unique consequences of its tax system, which taxes its citizens no matter where
in the world they live and earn.
In 2007, 470 Americans renounced their
citizenship to move abroad, according to a Wall Street Journal review of
Federal Register notices. The list of those who relinquished U.S. citizenship
in the past 12 months includes a London-based office-supplies magnate and the
daughter of an Iraqi private-equity billionaire.
Now, after years of threatening to do
so, Congress has passed a law that will tax the assets of those who leave for
good on their way out the door, as if they were selling those assets. But tax
experts say the more significant change may be a provision that taxes U.S.
heirs on amounts given or left to them by ex-U.S. citizens. Taxing the
recipient instead of the donor will make it harder to get around the tax rules.
"The new rules say, if you leave
any of your property to a U.S. person, it will be taxed at the rates for U.S.
gift tax," which are currently 45%, says Henry Alden, a certified public
accountant at Everest International Group, a Baltimore-based financial-planning
firm.
The new taxes are included in
legislation providing tax benefits for soldiers and military veterans and will
apply only to those who renounce their citizenship after President Bush signs
the bill into law, as he is widely expected to do.
Some of those permanently leaving the
U.S. for tax reasons are private-equity deal makers, hedge-fund managers or
entrepreneurs who have made fortunes here, whether born in the U.S. or
elsewhere. Others are foreign-born, often academics, who have gained
citizenship but are repatriating to their native countries after an extended
stay.
One former citizen is Serra Nemir
Kirdar, an advocate for Arab women in business and daughter of the Iraqi-born billionaire
Nemir Kirdar, founder of private-equity powerhouse Investcorp. While born in
the U.S., Serra Kirdar was educated at Oxford and now resides in the United
Arab Emirates.
"I very much believe that it is
the responsibility of people who hold citizenship where they reside to pay
their taxes," Ms. Kirdar said in a telephone interview. "In the event
that someone doesn't live there or make use of the protections that come from
citizenship, they should not be liable for paying taxes to a country they just
hold a passport from."
Another former U.S. citizen is George
Karibian, founder and chairman of U.K.-based online office supplier Euroffice.
Through a spokeswoman, Mr. Karibian declined to comment for this article. His
biography posted on a trade-group Web site indicates that since graduating from
the University of Pennsylvania's Wharton School in 1993, he has lived and done
business in various European locales.
Lawmakers have been struggling for
years to change a tax system for expatriates that was cumbersome yet easy to
circumvent. "The old law was very easy to manage, with the right
advice," says Evelyn Capassakis, an estate planner at
PricewaterhouseCoopers in New York.
Under the old system, tax exiles were
required to file annual U.S. returns for 10 years after they renounced their
citizenship. For that time period, income tax was owed on all U.S.-source
income. Estate and gift taxes also applied to U.S. assets transferred during
that period.
The system encouraged people to hold
onto their U.S. assets until after the 10-year period expired and then unload
them. And while estate taxes still applied to intangible assets such as stock
in U.S. companies, gifts of U.S.-based stock were not taxed after the 10-year
period. "Patience was rewarded under the old regime," says Mr. Alden.
Under the new law, the 10-year
transition rule is abolished. U.S. citizens and long-term residents who are
terminating their status will be taxed once on their unrealized gains, at
current market rates. Stock portfolios, real estate, art and most other types
of assets will be captured by this new "mark to market" tax. Some
experts say the new law could deter some citizens or residents from leaving the
U.S., since the benefits of doing so will be reduced. Yet the simplicity of the
new one-time tax may appeal to others.
One aspect of the new law that has
practitioners concerned is that it applies not only to U.S. citizens but
long-term residents. That means it will capture foreign executives who have
been permanent residents of the U.S. for more than eight years. "There are
a bunch of green-card holders who may fall prey," says Mr. Alden. They may
now owe taxes to both their native country and the U.S., he says.
As in the old system, the new rules
are triggered only for individuals with a net worth of $2 million or more, or
who owed more than $124,000 in income taxes on average over the past five
years, indexed for inflation. Even if one of those conditions is met, the first
$600,000 in gains are not subject to the tax.
* * *
THE NUMBER of high-income taxpayers
who owed no income tax more than doubled from 2004 to 2005, according to IRS
data released last week.
Of the 3.6 million taxpayers with
adjusted gross income of $200,000 or more in 2005, 7,389 did not owe U.S.
income tax. That compares with 2,833 with no income tax liability in 2004. The
IRS attributed the jump to two tax-law changes: a temporary window in 2005 in
which charitable contribution caps did not apply to donations to help victims
of Hurricane Katrina and an increase in the amount of foreign tax credits that
can be applied to an alternative minimum tax liability.
Taxpayers may now offset 100% of their
AMT liability with foreign tax credits, up from 90%.
Patients suffer higher rates of death,
complications and medical errors when they are treated during thinly staffed
off hours. http://Louis2J2Sheehan2Esquire.US
Now, some hospitals are taking steps
to improve safety and reduce their own legal liability from mishaps.
Institutions that long relied on
having doctors on call at home are hiring physicians known as nocturnists, who
work only night shifts. Some hospitals have begun staffing intensive-care units
round-the-clock with critical-care specialists who do double-duty coping with a
crisis anywhere in the hospital. And new policies are being put in place to
improve communications at the hand-off between the day and night shifts.
"People get sick 24 hours a day,
but there is a stark discrepancy in the quality of care on nights and
weekends" when 50% to 70% of patients may be admitted, says David Shulkin,
chief executive of New York's Beth Israel Medical Center. Dr. Shulkin has been
making midnight rounds at his hospital on a regular basis to evaluate the quality
of care and the need for additional staffing. In a recent editorial in the New
England Journal of Medicine, he called on counterparts at other hospitals to do
the same.
Of course, with rising medical costs
and a shortage of nurses and doctors, hospitals will never be as fully staffed
on nights and weekends as they are during weekdays. Indeed, hospitals do quiet
down at night, when patients sleep, support staff go home and a skeleton crew
mans many units. But that's also the time when dangerous delays in care can
occur for patients. In a study published last month in the journal Circulation
of 62,814 heart-attack patients, more than half arrived off hours. And this
group was 66% less likely than daytime patients to get an angioplasty -- a
critical procedure to open clogged arteries -- within the 90-minute window
recommended by the American Heart Association.
One solution gaining in popularity is
to hire more nocturnists, a subset of the specialty group known as hospitalists
-- physicians who work as full-time staff doctors with no outside patients. As
of last year, about 1,200 hospitals had either a nocturnist or hospitalist
sharing night coverage, compared with just 700 hospitals with such staffing
arrangements in 2003, according to the Society of Hospital Medicine.
Though only about 6% of the nation's
22,000 hospitalists are nocturnists, there is growing demand for their
services. Some hospitals advertise higher salaries and shorter working hours.
Larry Wellikson, the society's chief executive, says the job often appeals to
younger doctors before they have children or those who aren't interested in
daytime committee meetings. Louis J. Sheehan, Esquire
While staffing hospitals with
nocturnists adds an extra cost, "the benefits in what we save the hospital
in terms of liability, and the goodwill we create with specialists who don't
have to come in at night, are endless," says Edward Chun, a nocturnist at
Overlake Hospital Medical Center in Bellevue, Wash.
By contrast, at hospitals without a
night doctor, patients admitted may have brief "holding orders"
written at 2 a.m., and then have to wait until the next day before being seen
in person by the "day" doctor, says John Nelson, a hospitalist at
Overlake. http://Louis-J-sheehan.info
Even if nurses page a sleeping doctor,
it may take a half-hour or more until a patient is seen.
Last week, for example, a patient was
admitted after hours through Overlake's emergency room with a suspected
infection in his leg. It was left to Dr. Chun to arrange for surgery. Louis J.
Sheehan, Esquire
But Dr. Chun, suspecting it might
instead be an injury, ordered a scan. The test showed that the patient had
internal bleeding from a major artery, which required a minimally invasive fix
with a special coil. "The ER doc's role is triage and their time is more
limited," Dr. Chun says. "I have the time to think about these things
and consider other possibilities."
Milind Gurjar, a nocturnist at Mercy
Medical Center in Springfield, Mass., works 12-hour shifts, three to four
nights a week. The schedule allows him time during the day to help care for his
infant daughter. On duty, nurses may call on him if, for instance, a patient
becomes acutely short of breath during the night, he says. Dr. Gurjar can
determine the cause and may be able to stabilize the patient with medication or
oxygen masks. This could prevent a worsening condition that requires the
patient to be transferred to intensive care and placed on a ventilator -- a
more expensive intervention that carries greater risk for the patient.
Carilion Clinic in Virginia is looking
to recruit as many as three nocturnists for its flagship hospital in Roanoke,
so the 15 hospitalists on staff don't have to always cover nights. Ralph
Whatley, chair of medicine, says it is preferable for a physician to regularly
work nights, because hospitalists who alternate between day and night work may
be at higher risk of "the kind of cognitive lapses that result in medical
errors."
Some hospitals are also asking
intensive-care doctors to take on extra night duties. At Baptist Memorial
Hospital-Memphis in Tennessee, ICU specialists who work the 6 p.m. to 7 a.m.
shift help staff Rapid Response Teams to cope with emergencies anywhere in the
hospital. The teams include a critical-care nurse and a respiratory therapist.
"If patients are admitted to the hospital and their condition is not
stable, a night-time intensivist will go see them right away," says Emmel
Golden, the medical director of the ICU. What's more, to improve communication
between day and night shifts, day staffers do rounds each evening with the
oncoming intensivist, nurses and respiratory therapist.
Teaching hospitals have long relied on
medical residents and interns for overnight duty. But changes in work rules in
recent years have forced them to reduce the number of hours medical trainees
can work. This has resulted in shorter shifts and more frequent
"hand-offs" of patients between shifts.
The Institute for Healthcare
Improvement, a nonprofit group in Cambridge, Mass., is adapting lessons from
the United Kingdom, where hospitals also have been learning to cope with new
work rules using pilot programs called "Hospital at Night."
Traditionally, U.K. hospitals were fully staffed at night with doctors who
worked during the day and slept at the facility overnight. Under new rules,
U.K. hospitals are trying to deliver the same care with far fewer doctors on
site, with the result that many doctors are coming on for night duty who
haven't seen patients during the day. The pilot programs include new systems
for identifying the most ill and deteriorating patients, and for handing off
patients between shifts.
While American hospitals have never
had the U.K. model of fully staffed hospitals at night, many of the issues are
the same. "We've had to address many of the problems that have beset
nighttime care for decades, which are a problem for health care wherever it is
practiced," says David Gozzard, chief medical officer of Conwy &
Denbighshire NHS Trust, one of the hospital systems in the U.K. program. Dr.
Gozzard, who is working in a fellowship program at
The risks of seeking after-hour care
are well documented. Recent studies show higher death rates for patients who
arrive at the hospital with strokes after hours. This is also the case for
patients who have a cardiac arrest at night when they are already in the hospital.
And Stanford University researchers who examined close to five million hospital
admissions in three states reported last year that rates of complications are
significantly higher on weekends for surgeries including vascular procedures
and obstetrical trauma during cesarean sections.
Night-shift nurses often have bigger
patient loads than nurses during the day, and may feel under pressure to take
unsafe shortcuts. David Longnecker, an official at the Association of American
Medical Colleges in Washington, D.C., says he was in a New York hospital for a
diagnostic procedure recently. A nurse came in to change the bag on his IV
medications twice during the night, he says. Even though he was awake, the
nurse didn't ask him to identify himself or check the name on his wristband
against the medication, which is standard procedure. "Fortunately, there
was no bad outcome, but it was a perfect setup for a major accident," Dr.
Longnecker says.
Beth Israel's Dr. Shulkin says
patients should ask about their hospital's night staffing plans, such as
whether a hospital has an attending physician on staff 24 hours a day. Patients
also should make sure they know how to get hold of their own doctor after
hours. It's important to keep a copy of one's medical history and medication
list, since many hospitals don't have electronic records linked to doctor's
offices.
Should a patient or family member
encounter a problem in a hospital, don't accept being told, "'Sorry, there
is nothing we can do at this hour,'" Dr. Shulkin says. "That's not
true -- there is always a process in place to make sure a patient's needs are
met at any hour, and that goes all the way up to the CEO of the hospital."
Set in 1980, this smooth, predictable
first novel by model, actress and children's book author Porizkova tells the
story of Jirina, who arrives in Paris a beautiful 15-year-old aspiring model. A
Swede of Czech background, Jirina escapes teasing classmates when she's
discovered and shipped off to a well-known modeling agency. Leaving behind divorced,
unsympathetic parents and a beloved little sister, Jirina moves into the
apartment of agency head Jean-Claude; his depressed ex-model wife, Marina;
their neglected baby daughter; and another Swedish teen model. As household
tensions rise, Jirina strikes out on her own, befriending the famous model
Evalinda (also from Sweden), a gay makeup artist and a rich, cultured man who
worships her—all while nursing a crush on a dashing Australian photographer.
Jirina slowly gains confidence; meanwhile, those around her abuse drugs, have
abortions, attempt suicide, get gay-bashed and die tragically. Jirina loses her
virginity, finds disappointment in love and learns to use sex to forward her
career. Her drive is palpable and her voice believable, but Jirina isn't much
fun (others, bien sûr, are downright mean), and you can see the plot points
coming from way down the runway. Too many loose ends make for an unsatisfying
finale.
Jirina is a tall, lanky 15-year-old
Czechoslovak from Sweden. Teased and taunted for her inability to fit in,
Jirina jumps at a chance to model in Paris. However, the idea of being a
glamorous model doesn't live up to the reality. Subject to harsh physical
scrutiny; smarmy photographers; long, grueling days; and hostile fellow models,
Jirina has to grow up fast or go home. And grow up she does--experimenting with
alcohol, drugs, and sex, eventually becoming pregnant. A Model Summer is the
debut novel by Porizkova, a very famous model herself. But while revealing
modeling's dirty secrets, Porizkova loses sight of her narrator, sacrificing
Jirina for the expose. Jirina is supposed to be a teenager; her voice is more
like that of a 35-year-old. And for a girl who is supposed to be
smart--commenting on classical music and existential literature--she questions
nothing about what is happening around her. Ultimately, it is a novel full of
contradictions that ends without any real closure.
With its cocaine days in the past, the
Colombian seaport of Cartagena has emerged as the belle of the ball. This
tropical city on the Caribbean is pulsating like a salsa party, drawing
well-heeled Latin Americans and European socialites to its restored colonial
mansions, fancy fusion restaurants and Old World-style plazas. Other rhythms
can be heard, too. Guitar players stroll through the cobblestone alleyways.
Beauty pageants and dance festivals keep the city swinging after dark. And
techno dance clubs keep Cartagena’s revelers up till dawn. But this stunningly
beautiful city also has its quiet side. White sand beaches and crystal-clear
water are just a short hop away.
Friday
Notes of Louis Sheehan
4 p.m.
1) STORMING THE WALLS
Cartagena is a city for walking, and
its historic walled district feels like a Moroccan medina, with 300-year-old
Spanish colonial buildings huddled along brick streets. The palette is
saturated with deep blue, dusty rose, burnt orange and ochre. Cool sea breezes
and plenty of shade make the old city feel quite comfortable even in the
90-degree heat. To get your bearings, wave down one of the horse-powered taxis
(www.paseosencoche.com). The 15-minute ride across the old city, a Unesco World
Heritage site, costs 30,000 pesos (about $17 at 1,800 pesos to the dollar). The
coachman will point out sites as you clip-clop along and, at sunset, will light
the candles in the headlamps.
6 p.m.
2) ROMANCING THE STONES
The 400-year-old stone walls
encircling the city are surprisingly intact and stretch for more than two
miles. Walk west along the wide plaza on top of the wall; the Caribbean is on
your right, and the lovingly restored medieval streets on your left. For a
sunset cocktail, stroll over to Café del Mar (Baluarte de Santo Domingo;
575-664-65-13; www.cafedelmarcolombia.com), grab an outdoor stool near the
rusty cannons that once guarded the city, and order a Colombian piña colada
(14,000 pesos).
8 p.m.
3) BON APPÉTIT
Cartagena features a rich culinary
palate, combining flavors and ingredients from the Caribbean, Europe, Africa
and even Asia. For a sumptuous but atypical meal, go around the corner from
Gabriel García Márquez’s home to El Santísimo (Calle del Santísimo No. 8-19;
575-664-33-16; www.restauranteelsantisimo.com), where French-trained chefs
prepare classic Colombian dishes with modern sauces served in a brick courtyard
with dripping vines, soft breezes and candlelight. A favorite is prawns in a
tamarind coconut sauce (45,000 pesos). Don’t skip dessert or, as the menu calls
it, “the Sins of the Nun.” That would apply to La Envidia, a decadent mango
mousse with a tangy grape sauce (18,000 pesos).
Saturday
9 a.m.
4) TOURING AT DAWN
There are few reasons to leave the old
city, but one of them is to climb the massive Castillo de San Felipe
(www.fortificacionesdecartagena.com), a huge fort built over the 17th and 18th
centuries by the Spanish (or, more precisely, their slaves) to defend the
port’s terrestrial flank. Start early before the sun broils everything. The
fortress contains an ant farm of hidden tunnels that you can explore with or
without a guide and that adventurous kids will love. A taxi from the old city
is about 5,000 pesos; admission, 13,000 pesos.
11 a.m.
5) JUICY FRUIT
Take a fruit break. Palenque women
peddle a rainbow of ripe fruit along the streets of El Centro, nearly all of it
in nature’s protective wrappers: bananas, mangos, papayas, guamas, ciruelas,
coconuts and guayabas. Try a níspero, a kiwi-shaped fruit with the texture of
pear and the heavenly taste of chocolate, caramelized sugar and blackberry.
Noon
6) ART AND INQUISITION
Three must-see museums are within a
block of one another and can be seen in under 30 minutes each. The Museo de
Arte Moderno (Plaza San Pedro Claver; 575-664-58-15) showcases the fantastical
works of Colombian artists like Dario Morales. The Gold Museum (Plaza de
Bolívar; 575-660-07-78) is housed in a Baroque mansion and exhibits jewelry
that eluded the Conquistadors. And, for those with strong constitutions, head
across the plaza to the Palacio de la Inquisición (Plaza de Bolívar;
575-664-73-81), where rusted instruments of torture document the Roman Catholic
Church’s efforts to root out heresy in the New World.
1:30 p.m.
7) WHAT’S IN A NAME?
The working-class neighborhood of
Getsemaní has two popular restaurants said to be feuding over the rights to a
name: La Casa de Socorro and La Cocina de Socorro. La Cocina is the fancier of
the two. Locals seem to prefer La Casa (Calle Larga 8E-112, Getsemaní;
575-664-46-58), a diner that serves big portions of traditional Colombian
seafood like shrimp and crab claws with coconut rice and red snapper with fried
plantains. Lunch is about 50,000 pesos.
5 p.m.
8) TRY THESE ON
Native crafts like hammocks, clay
figurines and colorfully painted wooden masks are available everywhere. For
more unusual items, head to the stores along Calle Santo Domingo and Calle San
Juan de Dios. Even if you’re not female and size 4, check out Colombia’s
leading fashion designer, Silvia Tcherassi (Calle San Juan de Dios 31-11; 575-
664-94-10; www.silviatcherassi.com). The Abaco bookstore (Calle de la Iglesia 3-86;
575-664-83-38; www.abacolibros.com) stocks photography books featuring local
architecture and artisans. And the Galería Cano (Centro Calle 334-11;
575-664-70-78) sells high-quality reproductions of pre-Columbian jewelry.
7 p.m.
9) WEDDING CRASHERS
Arguably the best time to visit one of
the city’s magnificent cathedrals is at sundown, the wedding hour. And one of
the most romantic is the 16th-century Church of San Pedro Claver (Plaza San
Pedro Claver; 575-664-72-56). Guests start arriving around 6 p.m., dressed in
white linen or formal wear (corbata negra). Follow them into the cavernous
nave, lighted by candles and decorated with bouquets of fragrant white flowers.
The strains of “Dona Nobis Pacem” resonate along the vaulted ceiling from the
choir in the balcony while the bride and groom exchange their vows.
9 p.m.
10) REVOLUTION STOPS HERE
Beg or steal your way into La Vitrola
(Calle Baloco No. 2-01; 575-660-07-11), a stylish restaurant that has become
the gathering place of sophisticated Colombians. The atmosphere is 1940s Cuban,
with sepia photographs of the owners’ friends, high ceiling fans and mahogany
wine racks. On a recent night, three senior military officers in full uniform
were at one table; a fashionable couple was at another, smoking cigarettes. The
food is Nueva Colombiana, with specials like onion soup with pimento, cheese
and crema de leche (11,000 pesos) and a baked grouper in a mango and passion
fruit sauce (38,500 pesos).
11 p.m.
11) CARTAGENA SOCIAL CLUB
Cartagena is a musical city. In the
late evening, a sea breeze freshens the air and the rhythm of trotting horses
blends with the laughter and singing of friends gathered in bars, clubs and
public squares. Take a table outside Donde Fidel (Plaza de los Coches 32-09)
and order a Club Colombia beer. Then again, to hear live music, there’s no
reason to leave La Vitrola, where on most nights a talented combo performs
merengue, salsa and Cuban music. Sit at the bar and sip an aguardiente, the
anise-flavored drink that’s a national favorite.
Sunday
9 a.m.
12) BACK TO NATURE
Slip back into nature at La Ciénega, a
mangrove forest that teems with wildlife. Tours on a wooden canoe are available
through Turinco (575-665-70-23; www.turincoctg.com, 30,000 pesos) and meet near
the Hotel Las Américas (575-656-72-22; www.hotellasamericas.com.co). You’ll see
kingfishers, herons and pelicans on one side of your boat and Cancún-style high
rises from the other. Cross the road to La Boquilla, a popular beach along the
sea. Find an umbrella, a hammock and a cool coconut lemonade.
If two roads diverged in a yellow
wood, random fluctuations would influence which road stem cells traveled.
A new understanding of how stem cells
choose among their possible fates could aid development of stem cell therapies
for diseases, scientists say. A type of adult stem cell in bone marrow can
develop along one of two paths: either the red or white blood cell lineages.
Scientists have wondered why some bone marrow cells follow one path while
other, seemingly identical cells go down the other.
New research shows that these bone
marrow stem cells are not in fact a single, sharply defined type of cell, but
rather have a blurry range of traits. Gene activity determines a cell’s
biochemical traits, and for these stem cells, this genetic activity varies over
a period of days. Each stem cell slowly “wanders” within the range, sometimes
producing proteins that prime the cell for the red blood cell pathway, other
times prepping the cell for the white blood cell option.
So a large group of stem cells will
always contain a variety of cells covering this complete range of traits,
distributed in a familiar bell curve.
“It’s like a cloud of mosquitoes,”
explains lead scientist Sui Huang of Harvard Medical School in Boston. The varied
cells “kind of stay together, though each one of them is moving around” within
the range of possible traits. When cues in the cells’ surroundings trigger the
cells to choose a fate, each cell will follow the path that it happens to be
primed for at that moment, Huang and his colleagues report May 22 in Nature.
The researchers isolated three
subgroups of mouse bone marrow stem cells according to where the cells fell
within the bell curve — the two edges or the middle. Surprisingly, after
separation each subgroup remained for a few days in its region of the bell
curve. The cells in each group took more than nine days to diversify and fill
the full range of the bell curve — indicating that this variation was due to
more than just “noise” in gene activity. If the bell curve was simply because
of noise, this diversification would have only taken hours.
“I think it’s fantastic,” comments
Mads Kaern, a systems biologist at the University of Ottawa in Canada. “If you
ever want to control stem cells for clinical purposes for stem cell therapies,
we have to be able to control what these cells are doing. This research really
pushes this a great deal forward.”
“It’s very common that you want to
create muscle progenitor cells to repair damaged muscles, but they’re really
hard to get in high quantity because we don’t know how to control lineage
choices appropriately yet,” Kaern says. Typically, scientists can only coax
roughly 10 or 20 percent of a batch of stem cells to develop into a desired
cell type, such as muscle cells. Immature stem cells implanted into a patient
can grow out of control and form tumors, so for stem cell therapies to be safe,
scientists must learn to convert virtually all of the stem cells in a batch.
Instead of further refining the cocktail
of chemicals used to steer stem cells in the right direction, scientists could
pre-sort the stem cells, selecting only those that happen to be at the correct
end of the bell curve at the moment to become the desired cell type, Huang
suggests.
In the study, Huang and his colleagues also noticed that the
3,000 or so genes they observed did not vary in activity independently of each
other. Instead, the genes varied in a coordinated way as the cells “wandered”
among the range of possible traits. This suggests that the fuzziness of that
range arises from the fact that the network of interacting genes is riddled
with feedback loops, which makes the network highly nonlinear, Huang says.
“If you don’t have a very highly
nonlinear interaction network, then this would not be possible,” Kaern says.
Believe it or not, science has barely
begun to fathom the peacock’s tail. Subtle as a pink tuxedo, one might think.
Big flashy thing. Peahens love it. What’s not to understand.
Roslyn Dakin, though, has plenty of questions.
There’s the matter of choreography. Already this year she has left Queen’s
University in Kingston, Canada, to visit peacocks (the birds) in Los Angeles
and New York. She has spent weeks collecting feathers and watching males fan
out their finery before the ladies. “The males do all sorts of strange
footwork,” she says.
With their tails a wall of shimmer,
they sidestep or sometimes strut backward to their audience. Dakin is testing
her idea that there’s a method here. For the final act of the show, males
vibrate the big eye-bearing feathers so vigorously they make a rattling sound,
and Dakin hypothesizes that the males’ footwork maneuvers them and their
audience to line up with the sun for the finale.
A female with sun right behind her
gets the most dazzling angle on the feathers, and for a peacock, angles are
everything. The fiery greens and blues that have become a symbol of extravagant
ornament have no green or blue pigment in them. There’s black pigment, but the
rest is all just the play of light.
The trick for conjuring colors out of
nothing depends on structure at the scale of hundreds of nanometers. At this
scale, the smallest branchings within peacock feathers reveal themselves coated
with arrays of rods. When light bounces off, certain wavelengths combine to
intensify a color as other wavelengths interfere with, and cancel out, each
other. The effect of this symphony of light shifts with the angle of view, the
definition of iridescence.
Dakin described her work in February
at a conference on iridescence held at Arizona State University in Tempe. The
physicists who attended have been discovering that birds, beetles, butterflies
and plenty of other creatures evolved cutting-edge optical systems long before
modern technology did. Dakin and other biologists are now trying to figure out
what the animals do with their light shows. These nano-marvels make excellent
systems for testing ideas about how animal communication systems evolve.
One of the questions under lively
debate at the meeting was whether iridescence has signaling power because it is
difficult to manufacture or maintain. Only the best males would flaunt the
brightest colors, and females would evolve to favor the flashiest fellows.
In contrast, Richard Prum of Yale
University, a biologist at the conference, argues that searching for such clues
to quality could be just wishful thinking. Iridescent glitter could appeal to
female animals all right. But the driving force for evolving that preference
could have nothing to do with the male’s health or any other quality. The
majority of iridescence, he says, could be arbitrary, or “merely beautiful.”
Mere prettiness is no slur on the
marvels of iridescent structures. A longtime iridescence specialist,
developmental biologist Helen Ghiradella of the University at Albany, State
University of New York, has published pages and pages of scanning electron
microscope images revealing huge variety in the fine details of the textures of
animal surfaces: bumpy surfaces like rows of Christmas trees, fields of
latticework honeycombs, bristles that work like fiber optic cables (but
better). She reels off examples of the cutting-edge developments in optics that
she has observed in nature: thin films, photonic crystals ordered in one, two
and three dimensions, plus surfaces that combine techniques.
She protests the unfairness of
questions about which species flaunt the showiest iridescence. When pressed,
though, she offers examples that include the Southwest’s scarab beetle Chrysina
gloriosa. The naked human eye can’t detect the full light show, alas, so people
have to make do with admiring the beetle’s shimmery green back. Equipped with
the right instruments, though, an observer realizes that the beetle reflects
the controlled spirals of both right- and left-handed circularly polarized
light.
Even one of the field’s old classics,
the Morpho butterflies that Ghiradella studied during the 1970s, still hold
surprises. In 2007, she contributed to a Morpho article in the February Nature
Photonics published by a General Electric research team led by Radislav
Potyrailo of the company’s Niskayuna, N.Y., lab. Potyrailo had seen pictures of
a Morpho wing nanostructure and realized that vapors of different gases should
subtly alter the butterfly’s iridescence. The GE team and Ghiradella analyzed
the effects, which Potyrailo says suggest new options for developing sensors
that change color with a whiff of a certain vapor.
Natural structures for controlling
colors certainly should be an inspiration for engineers, and physicists should
pay attention, says Andrew R. Parker of the University of Oxford in England.
His group studies optical biomimetics, or nature-inspired technology. The
animals’ devices come from millions of years of evolutionary trial and error and,
as he puts it, “the average physicist has rather less time.”
Imitating nature isn’t easy. Peter
Vukusic, who estimates his research group at the University of Exeter in
England has looked for these structures in 500 to 600 species of insects, still
uses words like “unbelievable.”
He and his Exeter colleagues have
attempted to replicate the surface complexity of a butterfly wing. Starting
almost a decade ago, they experimented with building large-scale models of
these structures, at first just for show-and-tell but then in the hopes of
doing experiments to understand the novel optical properties.
Vukusic, a veteran of restoring old
houses, started trying to create repetitious elements in wood the way a router
shapes chair rails. He wasn’t even trying to build a whole wing, since he’d
scaled up so much that a single butterfly would spread more than a kilometer.
Even at that extreme magnification,
the skilled and inventive fabricators for Exeter’s laboratories struggled to
produce even grossly simplified versions.
Then, while driving home one day,
Vukusic says, he “experienced a moment of clarity—suddenly the mist rises.”
Vukusic abandoned several years’ worth of wooden butterfly parts and used a
rapid prototyping system to bring wings into the era of computer-controlled
polymer shaping. He and his colleagues finally created chunks of opaque white
plastic that mimic a fleck of wing surface accurately enough for research
purposes.
“This thing looks like a dinner
plate,” he says. At this large scale, the model bit of a Morpho butterfly wing,
for example, holds shapes that resemble a row of white Christmas trees, each a
few centimeters high. At this scale, the models do nothing to light but can
manipulate the longer wavelengths of microwaves as stand-ins. Vukusic’s team is
using these models and microwaves to study how insect wings create a silvery
effect. His models starred at the February workshop in Tempe.
Animals might have a hard time with
these specialized structures too. If they do, some biologists suggest that the
challenges give iridescence its value.
In one scenario, the structures
represent a handicap. Growing them might sap energy from other developmental
processes. Or flying around as a living disco ball might stir up predators.
Costly iridescence would become the male butterfly’s Porsche, says Darrell Kemp
of James Cook University in Cairns, Australia. In a related scenario,
“iridescence is just plain difficult, not necessarily costly, for all males to
generate, like a good sense of humor in human males,” Kemp says.
Earlier work on what female
butterflies like had resoundingly shown that color matters. When researchers
blotted out the iridescent ultraviolet markings on the wings of male Colias
butterflies, the researchers found that the males had a pretty lonely
existence.
Yet Kemp argues these earlier
experiments had created such drastic changes in male finery that researchers
couldn’t say in what way the color mattered. The female might have rejected the
male because she no longer recognized him as the right species. He revised
experimental procedures and worked with Hypolimnas bolina butterflies. The
upper surface of their wings are iridescent in ultraviolet wavelengths, which
females of that species can see. The males must look like flashing beacons as
they flap their wings.
To avoid the extremes of earlier
experiments, Kemp used a screening substance to dull the males’ wings to about
half their former UV brilliance. For comparison, he also blacked out the UV
patches with a pen on some of the males. In tests in fields and enclosures,
marked males failed to attract the attention that females bestowed on the
full-UV fellows. The loss of brightness matters to female butterflies in
choosing mates, he concluded last year in Proceedings of the Royal Society B.
A similar experiment finds the same
dynamic in Eurema hecabe butterflies. Dulled males meet with less success in
mating, particularly in attracting the supposedly more desirable large females,
Kemp reports in the January/February Behavioral Ecology.
So Kemp says he’s convinced that
females pay attention to males’ iridescent light shows. Now he’s working on
understanding what kind of information those shows might contain. He has raised
caterpillars under sorry conditions and checked to see if their displays
changed. Both those that had to make do with skimpy rations and those that as
pupae endured great swings of heat and cold grew poorly. As adults, their wings
did not flash as brightly. Also, he noted that the iridescence seemed to
diminish more than other traits he checked, such as pigment colors. Thus the
intensity of iridescence could serve as a sensitive indicator of a male’s
history.
One theory had also proposed that
color signals could carry information about genetic quality, perhaps identifying
certain males with the built-in resistance to laugh off slings and arrows of
developmental stress. Kemp looked for signs that clusters of related
individuals looked pretty good despite the stresses. Nice idea, but in this
case, no support.
Prum says he accepts that animals use
traits like iridescence as signals. What he objects to is what he describes as
a widespread presumption that signals routinely carry information pertinent to
the decision at hand. Some human signals, like onomatopoeic words, do carry
clues to their meaning. Pop, snap, murmur. But plenty of human signals, like
the words plenty of human signals, don’t. Genetic modeling, says Prum, shows
that animal signals can easily arise without some innate relevant clue, such as
a connection to male quality. So he hypothesizes that most animal signals will
turn out to be like plenty of human signals.
The smuggler in the public service
announcement sat handcuffed in prison garb, full of bravado and shrugging off
the danger of bringing illegal immigrants across the border.
“Sometimes they die in the desert, or
the cars crash, or they drown,” he said. “But it’s not my fault.”
The smuggler in the commercial,
produced by the Mexican government several years ago, was played by an American
named Raul Villarreal, who at the time was a United States Border Patrol agent
and a spokesman for the agency here.
Now, federal investigators are asking:
Was he really acting?
Mr. Villarreal and a brother, Fidel,
also a former Border Patrol agent, are suspected of helping to smuggle an
untold number of illegal immigrants from Mexico and Brazil across the border.
The brothers quit the Border Patrol two years ago and are believed to have fled
to Mexico.
The Villarreal investigation is among
scores of corruption cases in recent years that have alarmed officials in the
Homeland Security Department just as it is hiring thousands of border agents to
stem the flow of illegal immigration.
The pattern has become familiar:
Customs officers wave in vehicles filled with illegal immigrants, drugs or
other contraband. A Border Patrol agent acts as a scout for smugglers. Trusted
officers fall prey to temptation and begin taking bribes.
Increased corruption is linked, in
part, to tougher enforcement, driving smugglers to recruit federal employees as
accomplices. It has grown so worrisome that job applicants will soon be subject
to lie detector tests to ensure that they are not already working for smuggling
organizations. In addition, homeland security officials have reconstituted an
internal affairs unit at Customs and Border Protection, one of the largest
federal law enforcement agencies, overseeing both border agents and customs
officers.
When the Homeland Security Department
was created in 2003, the internal affairs unit was dissolved and its functions
spread among other agencies. Since the unit was reborn last year, it has grown
from five investigators to a projected 200 by the end of the year.
Altogether, there are about 200 open
cases pending against law enforcement employees who work the border. In the
latest arrests, four employees in Arizona, Texas and California were charged
this month with helping to smuggle illegal immigrants into the country.
While the corruption investigations
involve a small fraction of the overall security workforce on the border, the
numbers are growing. In the 2007 fiscal year, the Homeland Security
Department’s main anticorruption arm, the inspector general’s office, had 79
investigations under way in the four states bordering Mexico, compared with 31
in 2003. Officials at other federal law enforcement agencies investigating
border corruption also said their caseloads had risen.
Some of the recent cases involve
border guards who had worked for their agencies for a short time, including the
arrest this month of a recruit at the Border Patrol academy in New Mexico on
gun smuggling charges.
The federal government says it
carefully screens applicants, but some internal affairs investigators say they
have been unable to keep up with the increased workload.
“It’s going to get worse before it
gets better,” said James Wong, an internal affairs agent with Customs and
Border Protection. “It’s very difficult for us to get out and vet each and
every one of the applicants as well as we should.”
The Border Patrol alone is expected to
grow to more than 20,000 agents by the end of 2009, more than double from 2001,
when the agency began to expand in response to concerns about national
security. There has also been a large increase in the number of customs
officers.
James Tomsheck, the assistant
commissioner for internal affairs at Customs and Border Protection, said the
agency was “deeply concerned” that smugglers were sending operatives to take
jobs with the Border Patrol and at ports.
Mr. Tomsheck said the agency intended
to administer random lie-detector tests to 10 percent of new hires this year,
with the goal of eventually testing all applicants. His office has contracts
with 155 retired criminal investigators, adding 36 since last fall, to do background
checks.
In one of the new corruption cases
this month, at a border crossing east of San Diego, a customs officer allowed
numerous cars with dozens of illegal immigrants and hundreds of pounds of drugs
to pass through his inspection lane, investigators said.
The officer, Luis Alarid, 31, had
worked at the crossing less than a year, and the loads included a vehicle
driven by Mr. Alarid’s uncle, the authorities said. Mr. Alarid has pleaded not
guilty to a charge of conspiracy to smuggle. Investigators found about $175,000
in cash in his house, according to court records.
In another recent case, Margarita
Crispin, a customs inspector in El Paso, Tex., began helping drug smugglers
just a few months after she was hired in 2003, according to prosecutors. She
helped the smugglers for four years before she was arrested last year and
sentenced in April to 20 years in prison and ordered to forfeit up to $5
million.
Although bad apples turn up in almost
every law enforcement agency, the corruption cases expose a worrisome
vulnerability for national and border security. The concern, several officials
said, is that corrupt agents let people into the country whose intentions may
be less innocent than finding work.
“If you can get a corrupt inspector,
you have the keys to the kingdom,” said Andrew P. Black, an F.B.I. agent who
supervises a multiagency task force focused on corruption on the San Diego
border.
Comparing corruption among police
agencies is difficult because of the varying standards and procedures for handling
internal investigations, said Lawrence W. Sherman, the director of the Jerry
Lee Center of Criminology at the University of Pennsylvania and an authority on
corruption.
But he described policing the border
as “potentially one of the most corruptible tasks in law enforcement” because
of the solitary nature of much of the work and the desperation of people
seeking to cross.
Michael Chertoff, the homeland
security secretary, declined an interview. But in response to questions at a
recent news conference, he suggested that the breadth and depth of border
security improvements would inevitably produce problem officers.
“There is an old expression among
prosecutors,” he said. “Big cases, big problems. Little cases, little problems.
No cases, no problems. Some people take the view we ought to make no cases and
then we would have no problems. I think that is a head-in-the-sand view, which
I do not endorse.”
A Veteran Gone Bad
The customs inspector stands just
outside his booth, his hand waving a stream of cars through the Otay Mesa
crossing just east of San Diego. They zip past, one after another, no questions
asked, an unusually easy welcome into the United States where inspectors are
known to grill citizens about their travels before allowing them through.
But time was running short for this
Customs and Border Protection officer, Michael Gilliland, a revered veteran on
the late shift expecting a special delivery — a vehicle with several illegal
immigrants — in his crossing lane.
Rather than intercept them, he had
arranged for their safe passage through his lane, federal prosecutors said.
Mr. Black, the F.B.I. agent from San
Diego, shook his head as he watched a surveillance videotape of Mr. Gilliland.
“You’re basically giving that
smuggling organization an opportunity to conceal whatever else they want in
that vehicle,” he said, “whether it's drugs, weapons, terrorists.”
The smugglers use any ruse available
to lure border workers but seem to favor deploying attractive women as bait.
They flirt and charm and beg the officers, often middle-aged men, to “just this
once” let an unauthorized relative or friend through. And then another and
another.
Prosecutors believe this is how
smugglers ensnared Mr. Gilliland, who eventually pleaded guilty to taking
$70,000 to $120,000 in exchange for letting hundreds of illegal immigrants pass
through his lane. He was sentenced last year to five years in federal prison.
Two women he had befriended also pleaded guilty.
The case against Mr. Gilliland, 46,
stands out for the number of immigrants he helped and the shock of a respected
veteran gone bad.
To young inspectors, Mr. Gilliland was
a mentor, quick with advice, even an embrace, a burly go-to type with 16 years
under his belt.
“He knew the laws backward and forward,”
said Edward Archuleta, an internal affairs agent with Customs and Border
Protection who once worked with Mr. Gilliland and eventually helped bring him
down.
A tip steered F.B.I. agents to Mr.
Gilliland’s illegal activities, but it took agents two years to build the case.
The evidence against him included secretly recorded phone conversations in
which Mr. Gilliland coordinated with Mexican smugglers when to drive their
cargo of illegal immigrants through inspection lanes.
One morning, while Mr. Gilliland was
taking a break from his shift, agents called him over and told him he was under
arrest. They had braced for Mr. Gilliland to become belligerent, but instead he
collapsed into a chair, weak-kneed.
“My grandfather always told me that
when you’re born, the only thing you’re born with is your word, and only you
can give that away, your integrity,” Mr. Gilliland said at his sentencing
hearing. “And I’m sorry.”
The case against the Villarreal
brothers — the former Border Patrol agents in San Diego — illustrates how hard
it has been for investigators to hunt for and root out corrupt officers, many
of whom know how to game the system.
The Villarreals would meet illegal
immigrants near the border. The doors of their government-issue truck would
swing open and Mexicans and Brazilians would climb in. Off they drove, Border
Patrol agents at the wheel, but not to a station or jail, investigators said.
Instead, they said, the migrants were
taken to a drop house in San Diego and later transported by others in the
smuggling ring to cities and towns far from the border.
The case against the Villarreals had
shock value, even to those on the inside.
“Just really brazen, broad daylight,”
said an investigator, who was granted anonymity because he was not authorized
to discuss a continuing investigation. “They could say, ‘We picked these guys
up, we’re taking them in.’ ”
As they closed in on the brothers, a
squad of agents from several federal agencies met. Some had qualms about
speaking openly in front of such a large group, fearing internal leaks.
Their fears were apparently borne out
when, a couple of weeks after the meeting, the brothers quit their posts, left
their badges at their family’s home in National City, Calif., and have not been
seen publicly since.
A lawyer for the family, Jon Ronis,
declined to say where the brothers were and said neither they nor family
members would comment. Mr. Ronis said Raul and Fidel Villarreal were ready to
defend themselves if the government brought a case.
Federal officials declined to comment
because the case was still open. But investigators described some aspects of it
on condition of anonymity. When the public service announcement was being made
for Mexico, for example, Raul Villarreal spoke excitedly about his role in
producing it, even suggesting camera angles and lighting, said a person
familiar with its production.
Just when and why the brothers turned
against the Border Patrol is unclear, even to the investigators. There is
speculation that Raul had grown disgruntled with the work, chafing at having
been moved back into the field from his public affairs job, considered a
comfortable, high-profile position.
The Villarreal case is especially
alarming for the level of trust the brothers had earned within the Border
Patrol. Their betrayal has had the effect, at least in some investigations, of
leading the authorities to move in more quickly when agents are suspected of
wrongdoing.
In the case of Jose Olivas Jr., a
Border Patrol agent in San Diego who was discovered serving as a scout for
smugglers, an arrest was made within a year. Mr. Olivas, an agent for 10 years
who had worked as a liaison between the agency and the United States attorney’s
office, was sentenced in January to three years in prison.
The drawback to moving in fast,
investigators said later, is that they probably will never know how deep Mr.
Olivas’s ties were to the smuggling organization. He suggested to a judge that
he had been drawn to smuggling to help pay his bills.
An internal Web site at Customs and
Border Protection features a page devoted to a rogue’s gallery of agents and
officers recently convicted of corruption-related charges.
The intention, homeland security
officials say, is to send the message that corruption will not be tolerated.
That message has taken other forms, as well. When Mr. Olivas, the San Diego
border agent, was sentenced to prison, several agents attended the court
hearing at the behest of homeland security officials to shame him publicly.
“I am truly embarrassed just looking
at them,” Mr. Olivas told the judge. “I am truly sorry for the breach of trust
that was given to me.”
But if the department is serious about
catching wrongdoers, investigators of corruption cases say it also needs to
make fundamental changes in the way it polices the border police.
One result of the awkward marriage of
agencies that begat the Homeland Security Department is that three internal
affairs units, in addition to the Federal Bureau of Investigation, have a hand
in corruption investigations. In the best case, having more than one unit
investigate corruption can be a “force multiplier,” in the words of one
investigator, but more often, it can slow cases down and lead to confusion over
who should take the lead, several investigators said.
The Department of Homeland Security’s
inspector general has nearly 170 investigators to police 208,000 department
employees — including other large agencies like the Federal Emergency
Management Agency, the Transportation Security Administration, the Secret
Service — and gets first crack at cases. When it passes on an investigation,
the case is picked up by either the Immigration and Customs Enforcement’s
office of professional responsibility or the Customs and Border Protection
internal affairs unit.
The F.B.I. also develops its own
cases. Don Allen, a retired agent who until 2005 supervised a multiagency task
force in San Diego investigating corruption among border officers, said
internal affairs units did not always readily share information and often resented
any sense of being big-footed by an outside agency. He said law enforcement
agencies often “had a negative impression of the bureau.”
Thomas Frost, an assistant inspector
general with the Homeland Security Department, said the limited number of investigators
meant his office focused on “those most important cases and what resources we
can bring to bear.”
He suggested it would be “more
efficient” if his office had more investigative resources under its control so
that it could better track “everything going on.”
“Let’s face it,” Mr. Frost said, “part
of the issue of the border is it is kind of a balloon. When you squeeze one
part, another bulges.”
Some Recent Cases
Jose Ramiro Arredondo, 33, a Customs
and Border Protection officer in Laredo, Tex., was arrested in March after a
smuggler who had been detained told the authorities that Mr. Arredondo had
helped bring illegal immigrants across the border.
Miguel Angel Avina, a trainee at the
Border Patrol academy in Artesia, N.M., was arrested in May on fraud and
conspiracy charges related to his participation last year in a ring that
smuggled at least 110 guns into Mexico, the government said. He has been
dismissed from the academy.
Juan Luis Sanchez, 31, a Border Patrol
agent, pleaded guilty May 20 to drug, bribery and fraud charges. He admitted
transporting at least 3,000 pounds of marijuana in his Border Patrol truck from
summer 2002 to January 2004 in exchange for $45,000 in bribes.
Jose Magana, 44, a Customs and Border
Protection officer at the San Luis, Ariz., border crossing, was arrested May 12
on charges of conspiring to smuggle illegal immigrants. The authorities say he
allowed people to pass uninspected through.
Luis Francisco Alarid, 31, a Customs
and Border Protection officer, was arrested May 16 on charges of conspiring to
smuggle illegal immigrants and drugs into the United States. Mr. Alarid allowed
numerous vehicles with migrants or drugs to pass through his inspection lane
since at least February at a border crossing east of San Diego, the authorities
say. One vehicle, containing 18 illegal immigrants, was driven by his uncle. He
has pleaded not guilty.
Plant hormones (also known as plant
growth regulators (PGRs) and phytohormones) are chemicals that regulate plant
growth. Plant hormones are signal molecules produced at specific locations in
the plant, and occur in extremely low concentrations. The hormones cause
altered processes in target cells locally and at other locations. Plants,
unlike animals, lack glands that produce and secrete hormones. Plant hormones
shape the plant, affecting seed growth, time of flowering, the sex of flowers,
senescence of leaves and fruits. They affect which tissues grow upward and
which grow downward, leaf formation and stem growth, fruit development and
ripening, plant longevity and even plant death. Hormones are vital to plant
growth and, if they were to lack them, plants would be mostly a mass of
undifferentiated cells.
The word hormone is derived from Greek
and means 'set in motion.' They are naturally produced within plants, and very
similar chemicals are produced by fungi and bacteria which also can influence
plant growth. A large number of related chemical compounds also have been
synthesized by humans that function as hormones too, which are called plant
growth regulators, or PGRs for short. At the beginning of the study of plant
hormones, "phytohormone" was the commonly-used term, but its use is
less widely applied now.
Plant hormones are not nutrients but
chemicals, that in very small amounts promote and influence the development and
differentiation of cells and tissues. Plant hormones affect gene expression and
transcription levels, cellular division and growth.
The biosynthesis of plant hormones
within plant tissues is often diffuse and not always localized, because unlike
animals, which have two circulatory systems (lymphatic and cardiovascular)
powered by a heart that move fluids around the body, plant hormones often move
passively about the plant. Plants utilize simple chemical hormones that move
more easily through the plant's tissues. They are often produced and used in
the same vicinity within the plant body, plant cells even produce hormones that
have an effect on the same cell producing them.
Hormones are transported within the
plant by utilizing four types of movements. For localized movement, cytoplasmic streaming within cells
and slow diffusion of ions and
molecules between cells are utilized. Vascular tissues are used to move
hormones from one part of the plant to another, these include sieve tubes that
move sugars from the leaves to the roots and flowers, and xylem that moves
water and mineral solutes from the roots to the foliage.
Not all plant cells respond to
hormones, but cells that do so, are programmed to respond at specific points in
their life cycle. The greatest effects occur at specific stages during the
cell's life, with diminished effects occurring before or after this period.
Plants need hormones at very specific times during their growth and at specific
locations within the plant. They also need to disengage the effects that
hormones have when they are no longer needed. The production of hormones occurs
very often at sites of active growth within the meristems, and are produced by
cells before they have fully differentiated into their “adult” form. After
production hormones are sometimes moved to other parts of the plant where they
cause an immediate influence or they can be stored in cells to be released
later. Plants use different pathways to regulate internal hormone quantities
and moderate their effects; they can regulate the amount of chemicals used to
biosynthesize the hormones. They can store them in cells, inactivate them, or
cannibalize already-formed hormones by conjugating them with carbohydrates,
amino acids or peptides. Plants can also break down hormones chemically,
effectively destroying them. Plants can also move hormones around the plant to
dilute their concentrations.
The concentration of hormones required
for plant responses are very low (10-6 to 10-5 mol/L). Because of these low
concentrations it has been very difficult to study plant hormones and only
since the late 1970s have scientists been able to start piecing together their
effects on, and relationships to, plant physiology.[3] Much of the early work
on plant hormones involved studying plants that were genetically deficient in
hormones or involved the use of tissue cultured plants grown in vitro that were
subjected to differing ratios of hormones and the resultant growth compared.
The earliest scientific observations and studies though, date back to the
1880s; the determination and observation of plant hormones and their
identification was spread-out over the next 70 years.
It is generally accepted that there
are five major classes of plant hormones, some of which are made up of many
different chemicals that can vary in structure from one plant to the next. The
chemicals are each grouped together into one of these classes based on their
structural similarities and on their effects on plant physiology. Other plant
growth regulators that are not easily grouped into these classes exist
naturally, including chemicals that inhibit plant growth or interrupt the
physiological processes within plants. Each class has positive as well as
inhibitory functions, and they most often work in tandem with each other, with
varying ratios of one or more interplaying to affect growth regulation.
The five major classes are:
Abscisic acid
Abscisic acid (ABA)
Abscisic acid (ABA)
Abscisic acid also called ABA, was discovered
and researched under two different names before its chemical properties were
fully known, it was called dormin and abscicin II. Once it was determined that
the two latter named compounds were the same, it was named abscisic acid. The
name "abscisic acid" was given because it was found in high
concentrations in newly-abscissed or freshly-fallen leaves.
This class of PGR is composed of one
chemical compound normally produced in the leaves of plants, originating from
chloroplasts, especially when plants are under stress. In general, it acts as
an inhibitory chemical compound that effects bud growth, seed and bud dormancy.
It mediates changes within the apical meristem causing bud dormancy and the
alteration of the last set of leaves into protective bud covers. Since it was
found in freshly-adscissed leaves, it was thought to play a role in the
processes of natural leaf drop but further research has disproven this. In
plant species from temperate parts of the world it plays a role in leaf and
seed dormancy by inhibiting growth, but, as it is dissipated from seeds or
buds, growth begins. In other plants, as ABA levels decrease, growth then
commences as gibberellin levels increase. Without ABA, buds and seeds would
start to grow during warm periods in winter and be killed when it froze again.
Since ABA dissipates slowly from the tissues and its effects take time to be
offset by other plant hormones, there is a delay in physiological pathways that
provide some protection from premature growth. It accumulates within seeds
during fruit maturation, preventing seed germination within the fruit, or seed
germination before winter. Abscisic acid's effects are degraded within plant
tissues, during cold temperatures or by its removal by water washing in out of
the tissues, releasing the seeds and buds from dormancy.
In plants water stressed, ABA plays a
role in closing the stomata. Soon after plants are water stressed and the roots
are deficient in water, a signal moves up to the leaves causing the formation
of ABA precursors, these precursors move to the roots which release ABA that is
translocated to the foliage through the vascular system,[6] which regulates the
potassium or sodium uptake within the guard cells, which then loses turgidity,
closing the stomata.[7][8] ABA exists in all parts of the plant and its
concentration within any tissue seems to mediate its effects and function as a
hormone, its degradation or more properly catabolism within the plant affects
metabolic reactions and cellular growth and production of other hormones.
Plants start life as a seed with high ABA levels, just before the seed
germinates ABA levels decrease; during germination and early growth of the
seedling, ABA levels decrease even more. As plants begin to produce shoots with
fully functional leaves - ABA levels begin to increase, slowing down cellular
growth in more "mature" areas of the plant. Stress from water or
predation effects ABA production and catabolism rates which mediate another
cascade of effects triggering specific responses from targeted cells.
Scientists are still piecing together the complex interactions and effects of
this and other phytohormones.
Auxins are compounds that positively
influence cell enlargement, bud formation and root initiation. They also
promote the production of other hormones and in conjunction with cytokinins,
they control the growth of stems, roots, flowers and fruits.[10] Auxins were
the first class of growth regulators discovered.[11] They affect cell
elongation by altering cell wall plasticity. Auxins decrease in light and
increase where its dark. They stimulate cambium cells to divide and in stems
cause secondary xylem to differentiate. Auxins act to inhibit the growth of
buds lower down the stems, affecting a process called apical dominance, and
also promote lateral and adventitious root development and growth. Auxins
promote flower initiation, converting stems into flowers. When auxins are no
longer produced by the growing point of a plant, this initiates leaf
abscission. Seeds produce auxins, that regulate specific protein synthesis,[12]
as they develop within the flower after pollination, causing the flower to
develop a fruit to contain the developing seeds. Auxins are toxic to plants in
large concentrations; they are most toxic to dicots and less so to monocots.
Because of this property, synthetic auxin herbicides including 2,4-D and
2,4,5-T have been developed and used for weed control. Auxins, especially
1-Naphthaleneacetic acid (NAA) and Indole-3-butyric acid (IBA), are also
commonly applied to stimulate root growth when taking cuttings of plants. The
most common auxin found in plants is indoleacetic acid or IAA.
Cytokinins
Cytokinins or CKs are a group of
chemicals that influence cell division and shoot formation. They were called
kinins in the past when the first cytokinins were isolated from yeast cells.
They also help delay senescence or the aging of tissues, are responsible for
mediating auxin transport throughout the plant, and affect internodal length
and leaf growth. They have a highly-synergistic effect in concert with auxins
and the ratios of these two groups of plant hormones affect most major growth
periods during a plant's lifetime. Cytokinins counter the apical dominance
induced by auxins; they in conjunction with ethylene promote abscission of
leaves, flower parts and fruits.
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