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Appalachian Development Highway System
Map of the Appalachian Development Highway System
System information
Maintained by state or local governments
Length	3,090 mi (4,970 km)
Formed	March 9, 1965

ADHS signs for U.S. Route 78/Alabama State Route 4/ADHS Corridor X with their distinctive blue color. Most other states do not have distinctive highway shields for ADHS, however.

The Appalachian Development Highway System (ADHS) is a series of highway corridors in the Appalachia region of the eastern United States. The routes are designed as local and regional routes for improving economic development in the historically isolated region. It was established as part of the Appalachian Regional Development Act of 1965, and has been repeatedly supplemented by various federal and state legislative and regulatory actions. The system consists of a mixture of state, U.S., and Interstate routes. The routes are formally designated as "corridors" and assigned a letter. Signage of these corridors varies from place to place, but where signed are often done so with a distinctive blue-colored sign.

The Appalachian Regional Commission (ARC) forecast benefits of ADHS' completion by FY 2045 as the creation of 47,000 new jobs and $4.2 billion in gross regional product (GRP).

History

1966 map

In 1964, the President's Appalachian Regional Commission (PARC) reported to Congress that economic growth in Appalachia would not be possible until the region's isolation had been overcome. Because the cost of building highways through Appalachia's mountainous terrain was high, the region's local residents had never been served by adequate roads. The existing network of narrow, winding, two-lane roads, snaking through narrow stream valleys or over mountaintops, was slow to drive, unsafe, and in many places worn out. The nation's Interstate Highway System, though extensive through the region, was designed to serve cross-country traffic rather than local residents.

The PARC report and the Appalachian governors placed top priority on a modern highway system as the key to economic development. As a result, Congress authorized the construction of the Appalachian Development Highway System (ADHS) in the Appalachian Development Act of 1965. The ADHS was designed to generate economic development in previously isolated areas of the 13 Appalachian states, supplement the interstate system, and provide access to areas within the region as well as to markets in the rest of the nation.

The ADHS is currently authorized at 3,090 miles (4,970 km), including 65 miles (105 km) added in January 2004 by Public Law 108–199. A decade into construction, delays and cost increases mounted, attributed to:

• highway construction cost inflation
• upgraded construction guidelines following then-current Interstate Highway standards
• revised relocation assistance requirements
• delays associated with environmental protection, and
• Federal funding limitations

Periodic ADHS Completion Plan Reports were compiled to assess construction and the remaining cost-to-complete (C-to-C) forecasts, excerpts listed below.

Date	Open or constructing	Forecast C-to-C
1976	1,237 miles (1,991 km)	$7.9 B
1998	2,259 miles (3,636 km)	$8.5 B
2013	2,717 miles (4,373 km)	$11.4 B
2021	2,814 miles (4,529 km)	$10.3 B
2023	2,837 miles (4,566 km)	$9.7 B

By FY 2023, 2,837 miles (4,566 km)—approximately 91.8 percent of the authorized distance—were complete, open to traffic, or under construction. Many of the remaining miles will be among the most expensive to build. The ARC (the state governors) remain involved prioritizing, sequencing remaining corridor work. By 2040, 100% of ADHS' project miles are expected to be complete and open to traffic or, at least, partially complete.

Corridor Z across southern Georgia is not part of the official system, but has been assigned by the Georgia Department of Transportation.

Economic Results

Historically, highway investment has served as the basis for many US regional development policies and in 2008 the ADHS was deemed one of the more comprehensive programs to utilize the approach. To evaluate the effectiveness of such investments, land change modeling was used to compare 1976 "pre-" and 2002 "post-" highway conditions. The study focused on Ohio's US 32 portion of Corridor D and the 15 counties in close proximity; Adams, Athens, Brown, Clermont, Gallia, Highland, Hocking, Jackson, Meigs, Morgan, Pike, Ross, Scioto, Vinton, and Washington Counties. Using data acquired from the Landsat system of earth observational satellites, the comparison revealed slight, yet significant, levels of urban expansion within a 10 km (6.2 mile) band surrounding the new highway. Beyond this band land use was more stable, indicating even minor distance increases from the highway reduced the likelihood of further development. By 2016 new business growth along the corridor was drawing consumer traffic away from adjacent towns and county seats, causing revenue loss and unintended consequences for previously established town-centered businesses.

A 2016 economic assessment of ADHS' construction found regions of Appalachia benefitting differently. Case studies found some boosting tourism income, while others increased industrial activity or commercial/retail activity. Some regions had strong economic growth while others were dormant, the effects dependent on the pre-existing nature of the corridor, its population and workforce, its economic profile and proximity to surrounding business centers or markets. Case study excerpts from five corridors were:

• Corridor B (North Carolina and Tennessee). Though 305.5 miles (491.7 km) in total, this case study focused on the 88 miles (142 km) segment of Corridor B completed in 2003, passing through the Blue Ridge Mountains to connect western North Carolina with northeastern Tennessee. This project enabled improved access to the Port of Charleston and new residential and commercial developments near Weaverville, North Carolina. The highway led to a direct increase of around 4,600 new jobs in the area.

• Corridor D (Ohio & West Virginia). Its eastern 70-mile segment was completed in 1977, connecting interstates 77 and 79. The eastern segment shifted its economy from heavy industry into healthcare, education, government and education services. The highway also helped retain indigenous manufacturing activities and enabled around 1,000 new jobs.

• Corridor E (Maryland). Completion of this link (now part of I-68) enabled the region to boost its pre-existing tourism economy by drawing residents from Washington, DC and Baltimore; expand its pre-existing manufacturing and create new distribution (supply chain) activities. The highway project enabled an estimated 900 new jobs in the region.

• Corridor Q (Kentucky and Virginia). Improved connectivity through this mountainous region increased commuting range, facilitated commercial and retail development in several communities, development of an industrial park and a small business incubator. The highway had direct impact adding around 6,250 new jobs along the corridor.

• Corridor T (New York State). Becoming I-86 in 1998, this project saw economic gains with manufacturing jobs at several industrial parks, tourism jobs at a ski resort and a casino and service jobs at a call center. The corridor was critical in the location and expansion of manufacturing facilities for diesel engines, furniture and advanced ceramics – all needing interstate trucking connectivity. Over time, the highway was credited with generating over 3,200 jobs in the region.

A 2016–2019 study reported the cumulative ADHS construction efforts had led to economic net gains of $54 billion (approximately 0.4 percent of national income) and had boosted incomes in the Appalachian region by reducing the costs of trade. The 2021 ADHS Cost-to-Complete Estimate Report reiterated previous compilations that construction investments made between 1965–2015 contributed to the annual generation of over $19.6 B additional Appalachian business sales, representing $9+ B added GRP. Usage of the ADHS was saving 231 million hours of travel time annually, equivalent to a $10.7 B savings in transportation costs and worker productivity per year. The increased economic activity was helping to maintain or create over 168,000 jobs across the 13 Appalachian states. In 2021 ARC forecast that by 2045 ADHS' construction expenses would yield a return on investment (ROI) of 3.7, meaning $3.70 in benefits for every $1.00 invested in construction.

Employment gains credited to the ADHS were 16,270 new Appalachian jobs as of 1995; 42,190 by 2015.

List of ADHS corridors

Corridor A

Corridor A
Location	Sandy Springs, GA – Clyde, NC
Length	198.6 mi (319.6 km)

Corridor A is a highway in the states of Georgia and North Carolina. It travels from Interstate 285 (I-285) north of Atlanta northeasterly to I-40 near Clyde, North Carolina. I-40 continues easterly past Asheville, where it meets I-26 and Corridor B.

In Georgia, Corridor A travels along the State Route 400 (SR 400) freeway from I-285 to the SR 141 interchange southwest of Cumming. From here to Nelson, near the north end of I-575, Corridor A has not been constructed; its proposed path is near that of the cancelled Northern Arc. It begins again with a short piece of SR 372, becoming SR 515 when it meets I-575. SR 515 is a four-lane divided highway all the way to Blairsville. From Blairsville to North Carolina, the corridor has not been built, and SR 515 is a two-lane road.

The short North Carolina Highway 69 (NC 69) takes Corridor A north to U.S. Route 64 (US 64) near Hayesville. Corridor A turns east on US 64, and after some two-lane sections, it becomes a four-lane highway. Corridor A switches to US 23 near Franklin, and meets the east end of Corridor K near Sylva. From Sylva to its end at I-40 near Clyde, Corridor A uses the Great Smoky Mountains Expressway, which carries US 23 most of the way and US 74 for its entire length.

Corridor A-1

Corridor A-1
Location	Cumming, GA – Dawsonville, GA
Length	15.8 mi (25.4 km)

Corridor A-1 uses US 19/SR 400 from the point that Corridor A leaves it, at SR 141 near Cumming, northeast to SR 53 near Bright. SR 400 continues northeast as a four-lane highway from SR 53 to SR 60 south of Dahlonega; this section was built "with APL funds as a local access road".

Corridor B

Corridor B
Location	Asheville, NC – Lucasville, OH
Length	305.5 mi (491.7 km)

Corridor B is a highway in the states of North Carolina, Tennessee, Virginia, Kentucky, and Ohio. It generally follows U.S. Route 23 (US 23) from Interstate 26 (I-26) and I-40 near Asheville, North Carolina, north to Corridor C north of Portsmouth, Ohio.

Corridor B uses I-240 from its south end into downtown Asheville, where it uses US 23 (current and future Interstate 26) to Kingsport, Tennessee. The US 23 freeway ends at the Tennessee–Virginia state line, but US 23 is a four-lane divided highway through Virginia and into northeastern Kentucky.

At Greysbranch, Kentucky, Corridor B leaves US 23 to turn east on Kentucky Route 10 (KY 10) over the two-lane Jesse Stuart Memorial Bridge into Ohio. The short Ohio State Route 253 (OH 253) connects the bridge to US 52, a freeway that takes Corridor B north to Wheelersburg. US 52 continues west to Portsmouth, the proposed alignment of Corridor B continues north and northwest along Ohio State Route 823 to US 23 near Lucasville. The part of Corridor B north of SR 253 is also part of the I-73/74 North–South Corridor.

Corridor B-1

Corridor B-1
Location	Greenup, KY – Lucasville, OH
Length	18.0 mi (29.0 km)

Corridor B-1 travels from KY 10 to the north end of the Portsmouth Bypass. In Kentucky, it follows US 23 and US 23 Truck; after crossing the two-lane Carl Perkins Bridge into Ohio, it uses current and planned SR 852—a western bypass of Portsmouth—and US 23. Corridors B and B-1 both end near Lucasville, where Corridor C continues north along US 23 to Columbus.

Corridor C

Corridor C
Location	Lucasville, OH – Columbus, OH
Length	71.7 mi (115.4 km)

Corridor C is a highway in the U.S. state of Ohio. It is part of U.S. Route 23 (US 23), traveling from the north end of Corridor B near Lucasville north to Interstate 270 (I-270) south of Columbus. As of 2005, most of the road is a four-lane divided highway, but there are a few gaps yet to be built. Corridor C is part of the I-73/I-74 North–South Corridor.

Corridor C-1

Corridor C-1
Location	Jackson, OH – Chillicothe, OH
Length	27.3 mi (43.9 km)

Corridor C-1 is a connector from Corridor C near Chillicothe southeast to Corridor D near Jackson, Ohio, along US 35. It has been completed as a four-lane highway.

Corridor D

Main article: Corridor D

Corridor D
Location	Mount Carmel, OH – Clarksburg, WV
Length	232.9 mi (374.8 km)

Corridor D travels east–west from Interstate 275 (I-275), near Cincinnati, Ohio, to I-79, near Bridgeport, West Virginia. The corridor utilizes Ohio State Route 32 (SR 32) and U.S. Route 50 (US 50).

Decades after its completion Corridor D has provided mixed results with beneficial infrastructure, but with still unmet promises. Economic growth is evident in the corridor's western counties; several new hospitals, large car dealerships and several fast food restaurants were added. The Brown County Campus of Southern State Community College opened near Mount Orab, in a region where "there were no (previous) options for students, they had to drive an hour". The Mercy Health Mount Orab Medical Center and the Adams County Regional Medical Center were built alongside US 32. In 2006 a Southern Ohio Medical Center outreach branch opened in Adams County near the US 32 & SR-41 intersection at Peebles. Pike County's county seat, Jackson, has a developing retail thoroughfare running between US 32 and its historic downtown. But the corridor's advertised regional-prosperity never occurred. Counties along the corridor still have per capita median incomes below the state average and 20-35% below the national average and the gaps are not narrowing.

Radiation therapy#Side effects

Add Radiation Cystitis to side effects list. Also add topic of same name or merge with existing Hemorrhagic cystitis. See https://www.ncbi.nlm.nih.gov/books/NBK470594/ and https://pubmed.ncbi.nlm.nih.gov/20212517/ and https://emedicine.medscape.com/article/2055124-overview and https://my.clevelandclinic.org/health/diseases/22379-microhematuria

Myokymia

after updating, add to:

Category:Neurological disorders
Category:Extrapyramidal and movement disorders
Category:Radiation health effects

Article is badly out of date, focuses ONLY on Eyelid myokymia. Subject is much bigger. See https://emedicine.medscape.com/article/1141267-overview, https://emedicine.medscape.com/article/1213160-overview#a1, https://my.clevelandclinic.org/health/symptoms/17663-eye-twitching Two good definitions (an overview with facial or limb myofibers & technical) & canine refs at https://www.ncbi.nlm.nih.gov/pmc/articles/PMC8162615/ w/ differential for neuromyotonia

Crosslink with Radiation therapy#Late side effects

ref. https://www.ncbi.nlm.nih.gov/books/NBK482461/  Neuromyotonia is completed, has the NKB reference.

Several tie-ins to genetic mutations; ADCY5, KCNA1, KCNQ2. Inconsistent symptom descriptions for Myokymia. "peripheral nerve hyperexcitability (myokymia)" in https://www.ncbi.nlm.nih.gov/books/NBK32534/ " involuntary rippling movement of the muscles (myokymia)" in https://medlineplus.gov/download/ghr-summaries.xml "insuppressible and flickering movements confined to one eyelid (myokymia)" in https://www.ncbi.nlm.nih.gov/books/NBK542053/ "Superior oblique myokymia is ... characterized by spontaneous rhythmic contractions of the superior oblique muscle" at https://pubmed.ncbi.nlm.nih.gov/29279659/ with history ref to 1906.

causes

Focal platysma myokymia as the presenting symptom of cervical radiculopathy. Neurological sciences : official journal of the Italian Neurological Society and of the Italian Society of Clinical Neurophysiology [Neurol Sci] 2023 Jun; Vol. 44 (6), pp. 2221-2222. Date of Electronic Publication: 2023 Feb 27. Publisher: Springer-Verlag Italia Country of Publication: Italy NLM ID: 100959175 Publication Model: Print-Electronic Cited Medium: Internet ISSN: 1590-3478 (Electronic) Linking ISSN: 15901874 NLM ISO Abbreviation: Neurol Sci Subsets: MEDLINE

Limb Myokymia in Guillain-Barré Syndrome. Neurology India 68(1):p 230-233, Jan–Feb 2020. | DOI: 10.4103/0028-3886.279682. (Has Myo summary) (also see). Myokymia in Guillain-Barré syndrome. Neurology . 1983 Mar;33(3):374-6. doi: 10.1212/wnl.33.3.374. PMID: 6681885 DOI: 10.1212/wnl.33.3.374.

Post-Irradiation Facial Neuromyotonia/Myokymia: A Hemifacial Spasm Mimic. Abstract available. By: Swinnen BEKS; Koelman JHTM; van Rootselaar AF, Tremor and other hyperkinetic movements (New York, N.Y.) [Tremor Other Hyperkinet Mov (N Y)], ISSN: 2160-8288, 2021 Sep 20; Vol. 11, pp. 36; Publisher: Ubiquity Press; PMID: 34692229, Database: MEDLINE. Educational Value: These features are a red flag for (post-irradiation) facial neuromyotonia/myokymia which generally responds well to low dose carbamazepine.

Radiation induced subclinical brachial myokymia captured on muscle ultrasound. Clin Neurophysiol. 2020 May;131(5):1166-1167. doi: 10.1016/j.clinph.2020.02.001. Epub 2020 Feb 13. PMID: 32111545

Eyelid myokymia in patients with migraine taking topiramate. By: Medrano-Martínez V; Pérez-Sempere A; Moltó-Jordá JM; Fernández-Izquierdo S; Francés-Pont I; Mallada-Frechin J; Piqueras-Rodríguez L, Acta neurologica Scandinavica [Acta Neurol Scand], ISSN: 1600-0404, 2015 Aug; Vol. 132 (2), pp. 143-6; Publisher: Wiley-Blackwell; PMID: 25828425, Database: MEDLINE with Full Text.

Facial involvement in multiple sclerosis. Mult Scler Relat Disord. 2022 Nov:67:104110. doi: 10.1016/j.msard.2022.104110. Epub 2022 Aug 13. PMID: 35988397

Idiopathic generalized myokymia. Muscle & Nerve (journal). First published: January 1994 https://doi.org/10.1002/mus.880170106. https://onlinelibrary.wiley.com/doi/abs/10.1002/mus.880170106 SYMPTOMS /= TWITCHING.

Use of botulinum toxin type A for the treatment of radiation therapy-induced myokymia and neuromyotonia in a dog. J Am Vet Med Assoc. 2016 Mar 1;248(5):532-7. doi: 10.2460/javma.248.5.532. PMID: 26885596 (SYMPTOMS & TREATMENT)

chipmunk

connectivity between eastern chipmunk and chipmunk uncertain. Distinguish torpor vs hibernation in both.

Garter snake

Garter snake behavior addition- playing dead. https://thepetenthusiast.com/snakes-that-play-dead/

The related Apparent death. Needs article named Thanatosis. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC5769822/

Ewing's sarcoma

Add xrefs with Childhood cancer. Fits in section Malignancies of the musculoskeletal system

Ewing sarcoma#Treatment

side effects needs added; slow bone growth. SBG also impacts radiation therapy

https://www.cancer.org/cancer/types/ewing-tumor/treating/radiation-therapy.html

rare amputation example in "Acute and Long-Term Side-Effets" @ Springer (on phone homepage)

Disambiguation example to clean up Hygros' entry area

Eucalyptus

Eucalyptus (disambiguation)

Biology

collect Responsive encapsulations disussion, benefits here Responsive encapsulations have evolved in some species (ref location ???) to hygroscopically coordinate seed release with environmental conditions favorable to germination, e.g. Aizoaceae and Onoclea sensibilis. Hygroscopic coordination is also used in seed germination, as the normally impenetrable seed coat opens to wakening moisture aligned with favorable germination conditions, e.g. Trifolium pratense and Lupinus arboreus.

''Hygromorphic living materials for shape changing.'' Yao.

"A common example of a robust natural hygromorph is the pine cone". Reyssat.

collect future biomimetics and engineering hints "Active research" here -- -----

Also Hygromorphic materials for sustainable responsive architecture. by Holstov.

Additionally, functionally graded materials (FGMs), resulting from gradual changes in tissue structure and/or composition, which can produce repeatable, predictable hygrometric-controlled movement.

create a seed\spore distribution list

Format:MOS Wikipedia:Stand-alone lists

source info https://www.ecologycenter.us/plant-ecology/info-dnr.html Plant Nutrient Synchory Last Updated on Mon, 02 Jan 2023 | Plant Ecology

Vectors of Propagule Transport

The great number of different means of transport for propagules requires a structured overview of the vectors and mechanisms. Three large groups are distinguished:

• autochory, where the plant itself carries out the dispersal of its propagules,

• allochory, where the plant exploits different means of transport (vectors),

• atelochory, where dispersal is inhibited

Autochory

Plants have evolved many mechanisms which ensure distribution in different ways. In the simplest form of autochory, barochory, propagules are transported via gravity without special aids, a method that may only result in distribution over a wider area on steep slopes. Usually, the highest density of seeds is reached near the mother plant (seed shade). In contrast, with blastochory, propagules only find suitable growing sites by growth processes, e.g. via scions (vegetative shoots) or pedicels as in Cymbalaria muralis. In herpochory propagules move themselves short distances, e.g. the awns of Trifolium stellatum have hygroscopic characteristics, and twist with changes in humidity, dispersing the propagules (Fig. 4.2.1 B).

Special forms of autochory are summarised under the term ballochory, where propagules are propelled by a single impulse from the

■ Table 4.2.1. Distribution syndrome. Collected from Müller-Schneider (1977), Lindacher (1995), Frey and Lösch (1998)

Autochory (self dispersal) Barochory (transport via gravity)

Blastochory (dispersal via runners) Herpochory (transport via active creeping) Ballochory (self seeder)

- Zooballochory (impetus provided by animals)

- Anemoballochory (impetus provided by wind)

- Hydroballochory (impetus provided by water)

- Autoballochory (propulsion mechanisms based on sap pressure or drying)

Allochory (dispersal via a vector) Anemochory (transport via wind)

- Chamaechory (transport close to the soil because of large size and soil adherence to animals, etc.)

- Meteochory (transport in air for small seeds)

- Boleochory (transport started by wind, further dispersal assured by other mechanisms)

Hydrochory (transport in water)

- Nautochory (transport by movement in the sea)

- Bythisochory (drifting in flowing water)

- Ombrochory (transport via rain drops)

Zoochory (dispersal via animals), further differentiation by animal types, e.g.:

- Ornithochory, myrmecochory, etc.

- Epichory (transport by attachment of propagules to animals)

- Endochory (transport of propagules during passage through the gut)

- Stomatochory (transport in the mouth)

- Dysochory (transport of accidentally ingested propagules)

Hemerochory (dispersal by man)

- Ethelochory (deliberate dispersal)

- Speirochory (dispersal due to accidental dispersal with seeds)

- Agochory (unintentional dispersal)

Atelochory (transport and dispersal are not impeded)

mother plant. This may be triggered by animals, wind or even a raindrop. In autoballochory even such impulses are not required and the pressure is produced via differences in turgour (Ecbal-lium elaterium) or drying fruits or seed cases (Bauhinia purpurea, broom species). Seeds can be thrown over considerable distances, often several metres. Allochory Anemochory

Wind is a very important vector for allochoric seed transport. Even propagules blown by wind across the ground surface possess morphological adaptations. Such chamaechory is frequent in dry regions where there are rarely impediments to such transport. More important, however, is transport by air currents (meteochory). Small, light propagules which do not settle easily are transported in vertical - as well as horizontal -air streams across large distances, e.g. seeds of orchids or spores of cryptograms (so-called dust-fliers). Balloon fliers (Astragalus spinosus) possess special morphological adaptations, as do seeds or fruits with parachutes (Taraxacum officinale) and wings (Acer and Fraxinus species). Wind also triggers seed transport in plants which scatter seeds, e.g. the scatter mechanisms of Papaver species (boleochory). Hydrochory

Transport in, and with the help of, water is important for many plant species. Some propagules are able to float because of special tissues or large intercellular spaces, have a low specific weight and an external cover which is difficult to wet (Nymphea and Nuphar species). Coconuts, as well as seeds and seedlings of mangrove species, are even able to stay in salt water for long periods without damage. Seed dispersion in salt water and stagnant water is called nauto-chory; that in flowing water is called bythiso-chory. In flowing water seeds are transported by floating, and often drift over large distances. Small distances are often achieved by the impact of raindrops (ombrochory, e.g. Anastatica hiero-chuntica in dry regions). It is important that seeds in water do not germinate prematurely because of continuously moist conditions. Zoochory

The most important and ecologically most complex forms of allochoric distribution have devel oped in zoochory. Close interrelations between some plant and animal species point towards a long co-evolutionary development. This applies particularly to endochory, where propagules are taken up by the animal and pass through the intestines. During this time (retention time) they are transported by the animal. Most such seeds have a relatively hard shell and do not get damaged during passage through the gut, nor is it necessary for germination. However, after excretion, seeds are provided with very good starting conditions for germination in the nutrient-rich excrement. Some of the propagules are taken up randomly, particularly by large herbivorous mammals.

In addition to endochory, there are many other ways in which seeds are distributed by animals: Birds and bats, as well as ants, are specifically attracted; plants invest in nutrient-rich fruits and attract attention by striking colours or strong smells. Ants are particularly well known for their distribution processes (myrmecochory, however, this is a form of epichory, see below) and so are birds (ornithochory). The former carry seeds possessing lipid-rich elaiosomes (e.g. Corydalis cava) but the seeds remain untouched. This form of distribution is also called synchory. Birds react first and foremost to colours and are able to distinguish ripe seeds from unripe seeds.

In dysochory propagules are eaten and sometimes damaged. Such propagules, which are often starch-rich, are collected and accumulated as reserves (e.g. seeds of Pinus cembra collected by a bird, Nucifraga caryocatactes), but some pro-pagules still have a chance of surviving because not all hiding places are found again and some seeds are lost during transport. Some propagules are taken, together with the fruit pulp, into the mouth and spit out again (stomatochory) by many species of monkeys.

Propagules and their fruit pulp or endosperm are not always used as food. These means of attraction are often not developed at all, but still seeds or whole fruiting bodies are transported by animals. Propagules often possess glue-like excretions, glandular hair, barbs with awns and other outgrowths formed from the pericarp. Therefore, they are easily attached to fur and feathers and are transported until they fall off or are stripped off by animals during grooming and preening. In this form of transport, epicho-ry, animals distributing the seeds are not rewarded by food or energy and the plant invests less. In endochory close interaction exists be-

| Table 4.2.2. Relationship between propagules and dispersal by vertebrates. (Howe and Westley 1986, with additions)

Animal/animal group Propagule colour Propagule smell Propagule form Use to animals

Mammals In herds Birds In flocks

Brown

Green, brown

Little smell Without

Frugivor mammals in Yellow, green, white, Aromatic trees Bats

Ground living and frugivorous mammals

Frugivorous birds (obligate)

Frugivorous birds (facultative)

Furry or feathery orange, brown

Green, white, light Aromatic, musty yellow

Green, brown Without

Black, blue, red, green Without

Black, blue, red, white Without

Insignificant Without

Thick husked nuts, do not burst open

Seeds without wings and small nuts

Seeds often with arils, whole fruits, burst open

Diverse, often pendent fruit

Hard, over 50 mm long fruits, do not burst open

Big seeds with arils, whole seeds often burst open

Small seeds with arils, berries and stone fruits

Seeds

Seeds

Arils, pulp rich in proteins and sugars

Lipid- and starch-rich fruits

Lipid-rich fruits

Lipid- and proteln-rlch fruit flesh Mostly carbohydrate-rich fruit

Sticky and barbed hooks None tween plant and animals and distribution is regulated, sometimes even targeted. Epichory, in contrast, is often random. Some of the close links between characteristics of propagules and the animals distributing them are summarised in Table 4.2.2 (Howe and Westley 1986). Hemerochory

Man plays an increasingly important role in the recent history of plant distribution. In this particular allochoric form, hemerochory or anthro-pochory, any distances may be covered and all geographical and ecological barriers overcome. In ethelochory, plants are purposefully moved to different regions, e.g. to provide food or ornament. If distribution occurs unintentionally along with other propagules (e.g. weed seeds in seeds of cereal crops), this is called speirochory or in the case of random distribution agochory. Atelochory

The most important methods of distribution, autochory and allochory, are contrasted with atelochory (also called achory). This is a special form of distribution, as it is prevented. The consequence of this evolutionary development is that reproduction takes place at the site where the mother plant grows, which is favourable to the species. Examples are Arachis hypogaea or Trifolium subterraneum. After pollination pedicel and ovary penetrate into the ground.

Heterospory is widely used by therophytes in arid regions as the chance for survival are particularly good at sites where the mother plant is able to form fruits. Only some of the propagules are distributed to "conquer" new growing sites (Evenari et al. 1982).

TomStonehunter/sandbox
Conservation status
Secure (NatureServe)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Division:	Polypodiophyta
Class:	Polypodiopsida
Order:	Polypodiales
Suborder:	Aspleniineae
Family:	Onocleaceae
Genus:	Onoclea
Species:	O. sensibilis
Binomial name
Onoclea sensibilis L.

Onoclea sensibilis, the sensitive fern, also known as the bead fern, is a coarse-textured, medium to large-sized deciduous perennial fern. The name comes from its sensitivity to frost, the fronds dying quickly when first touched by it. It is sometimes treated as the only species in Onoclea, but some authors do not consider the genus monotypic.

Description

The sterile and fertile fronds of Onoclea sensibilis have independent stalks originating from the same rhizome, quite different from other ferns. The bright, yellow-green trophophylls (sterile fronds) are deeply pinnatifid and are typically borne at intervals along the creeping rhizome. The sterile fronds are deciduous with trophopods, swollen bases, that serve as over winter storage organs. The sterile fronds of O. var. sensibilis have a length of 1-1.3 m (3-4 ft) with 5-11 pinnae, leaf pairs, evenly spaced along the stipe. O. var. interrupta Maxim. fronds are shorter, 20-50 cm (8-20 in) long, with fewer pinnae, only 5-8 pairs.

Sporophyte "Beads"

The sporophylls (fertile fronds) are smaller, 20-45 cm (8-18 in) in length, non-green at maturity and have very narrow pinnae. They are persistent, standing 2–3 years. The sori comprise clusters of sporangia (spore cases) 2–4 mm (1/10-1/6 in) in diameter, like beads, on upright fertile fronds, hence the common name Bead fern.

Sori are typically bilaterally symmetrical, though leaf forms have been observed with pinnae fertile only on a single side of the rachis. This form, named O. sensibilis L. F. hemiphyllodes (Kiss & Kümmerle, 1926) and a second, O. sensibilis L. F. obtusilobata having pinnules flat (not curled or bead shaped), were deemed to be variations not meriting taxonomic recognition (J. M. Beitel et al. 1981).

Onoclea sensibilis' fiddleheads and first fronds

The fiddleheads have a pale reddish color.

Morphology

edits made large-scale hygromorphy The spermatogenesis process spans formation of spermatogenous cells to the release of spore. In homosporous ferns, like O. sensibilis L., developing spermatids are surrounded by two different walls at specific development stages, as opposed to a single wall reported in other species. Other differences include a delayed formation of the osmiophilic crest and during sperm release the cap cell removes intact, as opposed to forming a pore or collapsing altogether. Spores are monolete with the antheridium, or sporangium, containing either 32 or 64 sperm spores, usually being 64. Regardless of the number, the capsule's volume remains nearly the same.

The mechanics of spore release and its timing are controlled by springtime humidity. The small fertile margins, that in live-form held spore in tightly rolled structures, maintain their dry, leathery shape over winter. The pinnules' hygromorphic hygroscopic structures respond to spring's higher humidity by opening, releasing their spore into the air. Subsequent gametophytes are unisexual in early development, favoring cross-fertilization, later becoming bisexual to ensure species survival.

Taxonomy

Taxonomic History

WORK NEEDED

Kingdom: Plantae (Plants)

Subkingdom: Tracheobionta (Vascular plants)

Division: Pteridophyta (Ferns)

Class: Filicopsida

Order: Polypodiales

Family: Dryopteridaceae (Wood Fern family)

Genus: Onoclea Linnaeus (Sensitive fern)

Species: Onoclea sensibilis Linnaeus

IPNI (2021). International Plant Names Index (Royal Bot Gardens, KEW)

Kingdom: Plantae. Phylum: Filicinophyta. Class: Filicopsida. Order: Filicales. Family: Dryopteridaceae. Genus: Onoclea. by Nature Serve Explorer

Family: Woodsiaceae

1999 CRC World Dictionary of Medicinal and Poisonous Plants: Common Names ... By Umberto Quattrocchi Onocleaceae Aspleniaceae woodsaiceae

Onoclea orientalis, (Hook.) Hook., 1863, http://www.theplantlist.org/tpl1.1/record/tro-26624803

progression of tax history from L. onward: https://www.ncbi.nlm.nih.gov/pmc/articles/PMC3936591/

Aspleniineae has table example of dual tax threads

other north american family members 57 million years ago: https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/pteridaceae

Onocleaceae is here segregated, following recent molecular phylogenetic research (Smith & al. 2006). It was formerly often included in Dryopteridaceae or Woodsiaceae. https://alienplantsbelgium.myspecies.info/taxonomy/term/5329/descriptions

Sources matching with existing Taxonomy:

https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=17637#null

Species

The genus Onoclea was cast by Carl Linnaeus in 1751, separating from the fern's prior association with the Angiopteris genus. The binominal name, Onoclea sensibilis, was published in his 1753 Species Plantarum.

Varieties

Onoclea sensibilis has two geographically disjunctive varieties. Onoclea sensibilis var. sensibilis is native to North America; Canada's central and eastern regions and the United States' north, central and eastern regions. Onoclea sensibilis var. interrupta Maximowicz (aka Maxim.) is native to Southeast Siberia, Japan and China. The varietal difference is their ultimate height, O. var. interrupta Maxim. only reaching half the height of its sister.

Alternate names

Regional colloquial names for Onoclea sensibilis, the sensitive fern, focus on its characteristics.

• bead fern, an alternate name based on its fertile beaded pinnae (leaflets)
• bolletjesvaren; Dutch, meaning "ball fern"
• druebregne; Danish, meaning "grape fern"
• dwa’hũdes gananitsga’kwaʼ; Cayuga, meaning "deer, what they lie on"
• harilik pärljalg; Estonian, meaning "common pearl leg"
• helmisaniainen; Finnish, meaning "mother of pearl"
• pärlbräken; Swedish, meaning "pearl bracts"
• unì·suwεkwaʼ; Onondaga, meaning "bait"

Etymology

Onoclea sensibilis' name was descriptive. Onoclea comes from the Greek onos, meaning a vessel, and kleio, meaning to close, describing the closely rolled sori on its fertile fronds. Its species, from the Late Latin sensibilis, means sensitive, describing its high sensitivity to autumn's first frost and to drought.

Distribution and habitat

Onoclea sensibilis is native to Northern Hemisphere temperate regions; the Russian Far East, China and Eastern Asia, and a wide native distribution in Northern America. It ranges from Newfoundland south to Florida and west to Texas, the Rocky Mountains, North and South Dakota, Quebec, and Manitoba.

It has become naturalized in western Europe and New Zealand.

Onoclea sensibilis can be found at elevations from sea level up to 1,500 metres (4,900 ft) in fresh water habitats, not brackish, as it's spore germination ceases at NaCl levels ≥ 0.6%, moderately saline water and higher.

Onoclea sensibilis grows best in moist shaded or partially shaded areas, dwelling in a variety of swamp and wood habitats: wet meadows, thickets and bogs, as well as stream and riverbanks and roadside ditches. It tolerates extremely wet soils, appearing in soggy ground or at the very edge of water in shade or sun. The plant can tolerate dryer conditions in shade.

It prefers acidic (pH<6.8), loose, sandy to loam, limestone-based soils.

Ecology

Gametophyte (the flat thallus at the bottom of the picture) with a descendant sporophyte beginning to grow from it (the small frond at the top of the picture)

The Onoclea sensibilis plant has remained essentially unchanged over millions of years. A fifty-seven million year old fossil of Paleocene epoch flora shows specimens virtually identical to modern samples. It has a life cycle featuring alternation of generations, sexual and asexual reproduction; its sporophyte generation matures in autumn, casts its spores in the spring and the gametophyte generation follows. Surviving gametophytes require 5-10 years of growth before reaching their mature fern height.

Sensitive ferns propagate by both spore dispersion and rhizome growth. Its growth clusters attract local fauna where small wildlife find habitat, deer bed upon its dense mat and in winter wild turkeys use the fertile spore stalks as a secondary food source. They can become aggressive and a nuisance if established near preferable vegetation. The University of Maine's Cooperative Extension: Maine Wild Blueberries classifies the sensitive fern as a herbaceous broadleaf weed.

Its deciduous fronds do not tolerate freezing temperatures, however, the plant survives USDA hardiness zones 4-8, or minimum temperatures of -20 °C to -15 °C (-4 °F to 5 °F) having the Royal Horticultural Society's H6 rating. Winter survival is enhanced if the dried frond petiole bases are left intact.

Nutrient beneficial ectotrophic mycorrhizal associations may occur in Onoclea sensibilis, Pteridium aquilinum and Adiantum pedatum located in oak and hickory forests.

Onoclea sensibilis is a wetland indicator, listed as a Facultative Wetland Hydrophyte in the 2013 (US) National Wetland Plant List due to its observed affinity for wetter soils.

Opinion is mixed regarding the species' tolerance to disturbance of its growing environment. In one forest setting, a decade long decline was noticed following even single-cut tree felling operations. In other settings sensitive ferns appear opportunistic, disturbance not being a problem. They spread to form colonies, often the first species to inhabit disturbed areas.

Pests and diseases

Onoclea sensibilis hosts insects, fungi, bacteria and even a parasitic vine, Cuscuta gronovii (scaldweed), that can overgrow and constrict it.

Phlogophora iris- Olive angle shades, aka olive sausage

Insects

Insects feeding upon the Onoclea sensibilis target both its leaves and rhizome roots. Amphorophora ampullata fern aphids, Chirosia gleniensis fern miners and the larvae of sawflies Hemitaxonus dubitatus and Stromboceros delicatulus feed on its leaves. Larvae of moth species Phlogophora iris (olive angle shades, pictured), Callopistria cordata (silver-spotted fern moth) and Papaipema inquaesita (Sensitive Fern Borer) are known to feed on both stems and rhizomes.

Fungi

Parasitic fungi include Ceratobasidium anceps, causing frond and stem necrosis; Ceratobasidium cornigerum, covering stems with saprophyte growths; and Uredinopsis mirabilis, a distinct rust species unique to the Sensitive Fern. Invasive fungi like Taphrina filicina, Taphrina hiratsukae and Phyllactinia corylea, synonym Phyllactinia guttata can infect leaves, causing blisters or white powdery mildew.

Fungi can develop beneath beech trees, where aphid honeydew secretions accumulate; these strictly epiphyllous honeydew fungi, Sclerotiomyces colchicus and Scorias spongiosa (Schwein.) Fr., have been recorded on Onoclea sensibilis, where their sooty mold buildup impairs leaf function.

Bacteria

Onoclea sensibilis can host Burkholderia plantarii which causes stem lesions. B. plantarii is a pathogen of bacterial seedling blight in rice. In a multi-year study the weedy presence of O. sensibilis at rice paddy fields and a means to convey the bacterium (rainfall runoff) implicated it as the source of bacterial blight outbreaks when paired with enabling environmental conditions.

Toxicity

Pharmacology

Onoclea sensibilis has two internally-synthesized chemical defenses against insects. Ingesting any part of the plant introduces thiaminase enzymes and phytoecdysteroid hormones which can disrupt an insect's molting cycle, preventing its full development.

Mammal

Onoclea sensibilis has been implicated in equine poisoning and death, especially if eaten in quantity. The exact cause is unproven, but thiaminase poisoning, causing an extreme Vitamin B1 deficiency is suspected.

Human

Its human toxicity is not well defined; no specific warnings for Onoclea sensibilis have been found. Its summaries, however, frequently include precautionary statements that ferns, in general, may contain natural carcinogens and/or the enzyme thiaminase, the latter being dangerous in high concentration. Historically, some Native American peoples have consumed Onoclea sensibilis without apparent distress; see Food uses in this article.

Uses

Food

Onoclea sensibilis has limited value for food use, considered a famine food by some and reserved for times of scarcity. Cooking heat eliminates its thiaminase content. The Iroquois treated Onoclea sensibilis as an early springtime vegetable, prepared like spinach, the fiddleheads cooked and "seasoned with salt, pepper or butter" (Waugh, 1916). After removing the "brown scales" (sori), leaves were processed likewise. Its young shoots have been sold as delicacies in Asian markets.

Horticultural

It is cultivated as an ornamental plant in traditional and native plant gardens, and in natural landscaping and habitat restoration projects. It has gained the Royal Horticultural Society's Award of Garden Merit.Gardeners employ rhizome division and are aided by spore harvesting guides. Its decomposing fronds make an effective mulch, suppressing undergrowth. Plantings can become aggressive, weedy if not sited properly.

Decorative

Its cut fronds are used in dried flower arrangements.

Medicinal

Historically, Native American peoples used Onoclea sensibilis in both oral and topical forms for medicinal use. The Ojibwe people made a decoction of powdered, dried Onoclea sensibilis root to stimulate milk flow in female patients. The Iroquois did likewise and more extensively applied the root decoction for tuberculosis, fertility in women, pain and strength after childbirth, to treat and prevent baldness, as a gastrointestinal aid for swelling and cramps, for arthritis and infection. A fermented version of the decoction was used to start menses. A poultice of the top leaves was used for deep cuts and infection. A cold compound infusion of the entire fern plant was washed on sores and taken for venereal disease, e.g. gonorrhea.

Onoclea sensibilis has been found to contain the bioactive plant flavonols kaempferol and quercetin and has antioxidant properties in its var. interrupta's extracts.

An extract of Onoclea sensibilis has received future recommendation as an atherosclerosis treatment.

Links

Also see the Polish Wikipedia article for Onoclea sensibilis.

External links

• Boreal-forest.org
• Onoclea sensibilis. Web of Species: Biodiversity at Wellesley College and in New England.
• Onoclea sensibilis in L. Watson and M. J. Dallwitz (2004 onwards). The Ferns (Filicopsida) of the British Isles. delta-intkey.com
• Discussion of O. sensibilis' rhizome growth, branching and fern reproduction in A Phylogenetic Study of the Ferns of Burma

References

Category:Polypodiales Category:Ferns of Asia Category:Ferns of the Americas Category:Flora of China Category:Flora of Eastern Asia Category:Flora of the Russian Far East Category:Flora of Northern America Category:Ferns of the United States Category:Flora of Ontario Category:Garden plants of North America Category:Plants described in 1753 Category:Taxa named by Carl Linnaeus

Personal To Do List

Brachytherapy#Side effects#Long-term#Radiation

pmid=22949908 & Pradat in RILP intro

wiktionary.org/wiki/sensitive_fern

https://en.wiktionary.org/wiki/sensitive_fern has incorrect Etymology definition.

RIP- Radiation-induced fibrosis

https://europepmc.org/article/PMC/4573901?javascript_support=yes